Bivalvia | Unionida | Unionoidea | Margaritiferidae | Margaritiferinae

genus Margaritifera Schumacher, 1815: cladomics

7 Recent valid species (26 total species)

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 = 7 (26 total)
analyzed in 229 cladograms in 133 publications
maximum number of terminals in an analysis = 7 (mean = 2.24, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 113
maximum number of included terminals = 7, (mean = 3.5 median = 2)
proportion of cladograms that support basic monophyly = 0.96

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 Margaritifera cladogram distances
PCoA Ordination

NJ Tree of Margaritifera cladogram distances
NJ Tree

Cladograms

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

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
4 6 2 8 4 2 2 0 
  +-------- Margaritifera_margaritifera **  
  |                                      
  |  +----- Cumberlandia                 
--+  |                                   
  |  |  +-- Pseudunio                    
  |  |  |                                
  +--+  +-- Margaritifera_dahurica **       
     |  |                                
     |  +-- Margaritifera_middendorffi__1 **
     |  |                                
     |  +-- Margaritifera_middendorffi__2 **
     +--+                                
        +-- Margaritifera_falcata__1 **     
        |                                
        +-- Margaritifera_falcata__2 **     
        |                                
        +-- Margaritifera_falcata__3 **     
        |                                
        +-- Margaritifera_falcata__4 **

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
7 7 5 7 9 6 2 11 
                 +-- Cumberlandia               
  +--------------+                              
  |              +-- Pseudunio                  
  |                                             
--+              +-- Margaritifera_margaritifera **
  |  +-----------+                              
  |  |           +-- Margaritifera_dahurica **     
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_laevis **       
        +--+                                    
           |  +----- Margaritifera_middendorffi ** 
           +--+                                 
              |  +-- Margaritifera_marrianae **    
              +--+                              
                 +-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
7 7 5 7 625 4 2 11 
  +----------------- Cumberlandia               
  |                                             
--+              +-- Margaritifera_dahurica **     
  |  +-----------+                              
  |  |           +-- Margaritifera_margaritifera **
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_middendorffi ** 
        +--+                                    
           |  +----- Margaritifera_laevis **       
           +--+                                 
              |  +-- Margaritifera_hembeli **      
              +--+                              
                 +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 61 3 2 11 
              +-- Margaritifera_margaritifera **
  +-----------+                              
  |           +-- Margaritifera_dahurica **     
--+                                          
  |  +----------- Margaritifera_falcata **      
  +--+                                       
     |  +-------- Margaritifera_middendorffi ** 
     +--+                                    
        |  +----- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 191 4 2 11 
              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_dahurica **     
  |  +--------+                              
  |  |        +-- Margaritifera_margaritifera **
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     |  |                                    
     +--+     +-- Margaritifera_hembeli **      
        |  +--+                              
        |  |  +-- Margaritifera_marrianae **    
        +--+                                 
           |  +-- Margaritifera_laevis **       
           +--+                              
              +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
7 7 5 7 22 3 2 11 
                 +-- Cumberlandia               
  +--------------+                              
  |              +-- Pseudunio                  
  |                                             
--+              +-- Margaritifera_margaritifera **
  |  +-----------+                              
  |  |           +-- Margaritifera_dahurica **     
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_laevis **       
        +--+                                    
           |  +----- Margaritifera_hembeli **      
           +--+                                 
              |  +-- Margaritifera_middendorffi ** 
              +--+                              
                 +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 3 7 193 4 2 10 
  +-------------- Cumberlandia               
  |                                          
--+  +----------- Pseudunio                  
  |  |                                       
  |  |  +-------- Margaritifera_falcata **      
  +--+  |                                    
     |  |     +-- Margaritifera_dahurica **     
     |  +-----+                              
     +--+     +-- Margaritifera_margaritifera **
        |                                    
        |  +----- Margaritifera_laevis **       
        |  |                                 
        +--+----- Margaritifera_middendorffi ** 
           |                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 27 3 2 11 
              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_dahurica **     
  |  +--------+                              
  |  |        +-- Margaritifera_margaritifera **
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     |  |                                    
     +--+     +-- Margaritifera_hembeli **      
        |  +--+                              
        |  |  +-- Margaritifera_marrianae **    
        +--+                                 
           |  +-- Margaritifera_laevis **       
           +--+                              
              +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
7 7 5 7 21 4 2 11 
              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_dahurica **     
  |  +--------+                              
  |  |        +-- Margaritifera_margaritifera **
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     |  |                                    
     +--+     +-- Margaritifera_middendorffi ** 
        |  +--+                              
        |  |  +-- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 21 4 2 11 
                 +-- Cumberlandia               
  +--------------+                              
  |              +-- Pseudunio                  
  |                                             
--+              +-- Margaritifera_margaritifera **
  |  +-----------+                              
  |  |           +-- Margaritifera_dahurica **     
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_laevis **       
        +--+                                    
           |  +----- Margaritifera_middendorffi ** 
           +--+                                 
              |  +-- Margaritifera_hembeli **      
              +--+                              
                 +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 61 3 2 11 
              +-- Margaritifera_dahurica **     
  +-----------+                              
  |           +-- Margaritifera_margaritifera **
--+                                          
  |  +----------- Margaritifera_falcata **      
  +--+                                       
     |  +-------- Margaritifera_middendorffi ** 
     +--+                                    
        |  +----- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 6 3 2 11 
              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_dahurica **     
  |  +--------+                              
  |  |        +-- Margaritifera_margaritifera **
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     |  |                                    
     +--+     +-- Margaritifera_middendorffi ** 
        |  +--+                              
        |  |  +-- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_marrianae **    
           +--+                              
              +-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
7 7 5 7 19 4 2 11 
  +----------------- Cumberlandia               
  |                                             
--+              +-- Margaritifera_dahurica **     
  |  +-----------+                              
  |  |           +-- Margaritifera_margaritifera **
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_middendorffi ** 
        +--+                                    
           |  +----- Margaritifera_laevis **       
           +--+                                 
              |  +-- Margaritifera_hembeli **      
              +--+                              
                 +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 27 3 2 11 
           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  |  |                                    
  +--+     +-- Margaritifera_hembeli **      
     |  +--+                              
     |  |  +-- Margaritifera_marrianae **    
     +--+                                 
        |  +-- Margaritifera_laevis **       
        +--+                              
           +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
7 7 4 7 193 4 2 11 
  +----------- Cumberlandia               
  |                                       
  +----------- Pseudunio                  
  |                                       
--+        +-- Margaritifera_dahurica **     
  |  +-----+                              
  |  |     +-- Margaritifera_margaritifera **
  |  |                                    
  +--+  +----- Margaritifera_laevis **       
     |  |                                 
     |  |  +-- Margaritifera_falcata **      
     +--+--+                              
        |  +-- Margaritifera_middendorffi ** 
        |                                 
        |  +-- Margaritifera_hembeli **      
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 134 4 2 11 
  +-------------------- Cumberlandia               
  |                                                
--+  +----------------- Pseudunio                  
  |  |                                             
  +--+              +-- Margaritifera_dahurica **     
     |  +-----------+                              
     |  |           +-- Margaritifera_margaritifera **
     +--+                                          
        |  +----------- Margaritifera_falcata **      
        +--+                                       
           |  +-------- Margaritifera_laevis **       
           +--+                                    
              |  +----- Margaritifera_middendorffi ** 
              +--+                                 
                 |  +-- Margaritifera_hembeli **      
                 +--+                              
                    +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 61 3 2 11 
           +-- Margaritifera_margaritifera **
  +--------+                              
  |        +-- Margaritifera_dahurica **     
--+                                       
  |  +-------- Margaritifera_falcata **      
  |  |                                    
  +--+     +-- Margaritifera_laevis **       
     |  +--+                              
     |  |  +-- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_hembeli **      
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
7 7 5 7 4 3 2 11 
           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  |  |                                    
  +--+     +-- Margaritifera_laevis **       
     |  +--+                              
     |  |  +-- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_marrianae **    
        +--+                              
           +-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
6 6 4 6 24 3 2 11 
           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_laevis **       
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 6 4 6 132 3 2 11 
           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_laevis **       
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 6 4 6 7 4 2 11 
              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_margaritifera **
  |  +--------+                              
  |  |        +-- Margaritifera_dahurica **     
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     +--+                                    
        |  +----- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_middendorffi ** 
           +--+                              
              +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 6 4 6 132 3 2 11 
           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_laevis **       
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 6 4 6 24 3 2 11 
           +-- Margaritifera_margaritifera **
  +--------+                              
  |        +-- Margaritifera_dahurica **     
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_laevis **       
     +--+                                 
        |  +-- Margaritifera_marrianae **    
        +--+                              
           +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
6 6 4 6 70 3 2 11 
  +-------------- Pseudunio                  
  |                                          
--+           +-- Margaritifera_margaritifera **
  |  +--------+                              
  |  |        +-- Margaritifera_dahurica **     
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     +--+                                    
        |  +----- Margaritifera_middendorffi ** 
        +--+                                 
           |  +-- Margaritifera_laevis **       
           +--+                              
              +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 8 5 7 3 2 2 11 
  +----------- Margaritifera_margaritifera **
  |                                       
  |     +----- Margaritifera_dahurica__1 **  
  |     |                                 
  +-----+----- Margaritifera_dahurica__2 **  
--+     |                                 
  |     |  +-- Pseudunio                  
  |     +--+                              
  |        +-- Cumberlandia               
  |                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_laevis **       
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 6 4 6 4 2 2 11 
  +----------------- Pseudunio_auricularius__1  
  |                                             
  +----------------- Pseudunio_auricularius__2  
--+                                             
  |  +-------------- Cumberlandia               
  |  |                                          
  +--+           +-- Margaritifera_margaritifera **
     |  +--------+                              
     |  |        +-- Margaritifera_dahurica **     
     +--+                                       
        |  +-------- Margaritifera_falcata **      
        +--+                                    
           |  +----- Margaritifera_laevis **       
           +--+                                 
              |  +-- Margaritifera_middendorffi ** 
              +--+                              
                 +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 6 4 6 70 3 2 11 
           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_marrianae **    
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
5 5 3 5 7 2 2 10 
           +-- Pseudunio                  
  +--------+                              
  |        +-- Cumberlandia               
  |                                       
--+        +-- Margaritifera_dahurica **     
  |  +-----+                              
  |  |     +-- Margaritifera_margaritifera **
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
5 5 3 5 4 3 2 10 
  +-------------- Cumberlandia               
  |                                          
--+  +----------- Pseudunio                  
  |  |                                       
  +--+        +-- Margaritifera_dahurica **     
     |  +-----+                              
     |  |     +-- Margaritifera_margaritifera **
     +--+                                    
        |  +----- Margaritifera_falcata **      
        +--+                                 
           |  +-- Margaritifera_laevis **       
           +--+                              
              +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
5 5 3 5 5 3 2 10 
  +----------------- Pseudunio_marocanus        
  |                                             
--+  +-------------- Pseudunio_auricularius     
  |  |                                          
  +--+  +----------- Cumberlandia               
     |  |                                       
     +--+        +-- Margaritifera_dahurica **     
        |  +-----+                              
        |  |     +-- Margaritifera_margaritifera **
        +--+                                    
           |  +----- Margaritifera_falcata **      
           +--+                                 
              |  +-- Margaritifera_middendorffi ** 
              +--+                              
                 +-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
5 7 3 7 7 2 2 9 
  +-------- Margaritifera_dahurica **        
  |                                       
  +-------- Margaritifera_margaritifera__1 **
  |                                       
  +-------- Margaritifera_margaritifera__2 **
  |                                       
--+-------- Margaritifera_margaritifera__3 **
  |                                       
  |     +-- Pseudunio                     
  +-----+                                 
  |     +-- Cumberlandia                  
  |                                       
  |  +----- Margaritifera_falcata **         
  +--+                                    
     |  +-- Margaritifera_middendorffi **    
     +--+                                 
        +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
5 5 3 5 6 2 2 10 
  +----------- Pseudunio                  
  |                                       
  +----------- Cumberlandia               
--+                                       
  |        +-- Margaritifera_dahurica **     
  |  +-----+                              
  |  |     +-- Margaritifera_margaritifera **
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
5 5 3 5 4 3 2 10 
  +-------------- Cumberlandia               
  |                                          
--+  +----------- Pseudunio                  
  |  |                                       
  +--+        +-- Margaritifera_dahurica **     
     |  +-----+                              
     |  |     +-- Margaritifera_margaritifera **
     +--+                                    
        |  +----- Margaritifera_falcata **      
        +--+                                 
           |  +-- Margaritifera_middendorffi ** 
           +--+                              
              +-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
4 4 2 4 32 2 2 9 
  +----------- Cumberlandia               
  |                                       
--+  +-------- Pseudunio                  
  |  |                                    
  +--+     +-- Margaritifera_margaritifera **
     |  +--+                              
     |  |  +-- Margaritifera_dahurica **     
     +--+                                 
        |  +-- Margaritifera_falcata **      
        +--+                              
           +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
4 4 0 4 32 0 0 9 
  +-- Margaritifera_dahurica **     
  |                              
  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_laevis **       
  |                              
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
4 4 2 4 87 2 2 10 
        +-- Cumberlandia               
  +-----+                              
  |     +-- Pseudunio                  
  |                                    
--+     +-- Margaritifera_middendorffi ** 
  |  +--+                              
  |  |  +-- Margaritifera_laevis **       
  +--+                                 
     |  +-- Margaritifera_dahurica **     
     +--+                              
        +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
4 4 2 4 4 3 2 9 
  +----------- Cumberlandia               
  |                                       
--+  +-------- Pseudunio                  
  |  |                                    
  +--+     +-- Margaritifera_margaritifera **
     |  +--+                              
     |  |  +-- Margaritifera_dahurica **     
     +--+                                 
        |  +-- Margaritifera_falcata **      
        +--+                              
           +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
4 4 2 4 87 2 2 10 
        +-- Pseudunio                  
  +-----+                              
  |     +-- Cumberlandia               
  |                                    
--+     +-- Margaritifera_middendorffi ** 
  |  +--+                              
  |  |  +-- Margaritifera_laevis **       
  +--+                                 
     |  +-- Margaritifera_dahurica **     
     +--+                              
        +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
4 5 0 4 113 0 0 10 
  +-- Gibbosula                  
  |                              
  +-- Margaritifera_dahurica **     
  |                              
--+-- Margaritifera_margaritifera **
  |                              
  +-- Margaritifera_laevis **       
  |                              
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
3 3 1 3 7 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
3 3 1 3 66 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
3 3 1 3 13 0 0 9 
  +----- Margaritifera_margaritifera **
--+                                 
  |  +-- Margaritifera_falcata **      
  +--+                              
     +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
3 3 1 3 93 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 71 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
3 3 1 3 0 0 0 9 
  +----- Margaritifera_margaritifera **
--+                                 
  |  +-- Margaritifera_falcata **      
  +--+                              
     +-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
3 3 1 3 71 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 53 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
3 3 1 3 93 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 39 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
3 3 1 3 71 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
3 3 1 3 38 3 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 85 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 35 1 1 9 
  +-------- Cumberlandia               
--+                                    
  |  +----- Margaritifera_margaritifera **
  +--+                                 
     |  +-- Margaritifera_falcata **      
     +--+                              
        +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
3 3 1 3 83 2 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 38 3 2 9 
  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 36 1 1 9 
  +-------- Cumberlandia               
--+                                    
  |  +----- Margaritifera_falcata **      
  +--+                                 
     |  +-- Margaritifera_marrianae **    
     +--+                              
        +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 20 1 1 -- 
  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 12 0 0 -- 
  +-- Margaritifera_middendorffi **
--+                             
  +-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
2 2 0 2 9 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 42 1 1 -- 
  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_dahurica **
  +--+                         
     +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 14 0 0 -- 
  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 12 0 0 -- 
  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 10 0 0 -- 
  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Margaritifera_falcata ** 
--+                         
  +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 2 0 2 4 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 42 1 1 -- 
  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_dahurica **
  +--+                         
     +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 55 0 0 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
2 2 0 2 120 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 12 0 0 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 44 1 1 -- 
  +----- Cumberlandia          
--+                            
  |  +-- Margaritifera_dahurica **
  +--+                         
     +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 74 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 4 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 30 1 1 -- 
  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 47 1 1 -- 
  +----- Cumberlandia          
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 2 0 2 17 1 1 -- 
  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 6 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 4 19 0 0 -- 
  +-------- Margaritifera_laevis__2 **   
--+                                   
  |  +----- Margaritifera_laevis__1 **   
  +--+                                
     |  +-- Margaritifera_middendorffi **
     +--+                             
        +-- Margaritifera_laevis__3 **

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

iTtTbiriSoTroTroSiD
2 2 0 2 10 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 3 12 1 1 -- 
  +----- Cumberlandia               
  |                                 
--+  +-- Margaritifera_margaritifera **
  |  |                              
  +--+-- Margaritifera_dahurica__1 **  
     |                              
     +-- Margaritifera_dahurica__2 **

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

iTtTbiriSoTroTroSiD
2 2 0 2 20 1 1 -- 
  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 37 0 0 -- 
  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 31 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 5 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 6 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 26 1 1 -- 
  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 36 1 1 -- 
  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 2 0 2 17 0 0 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 2 0 2 4 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 46 1 1 -- 
  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 2 0 2 8 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 38 1 1 -- 
  +----- Cumberlandia          
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 15 0 3 14 0 0 20 
  +-------------------- Margaritifera_margaritifera__2 **
  |                                                   
  |                 +-- Margaritifera_margaritifera__1 **
  |  +--------------+                                 
--+  |              +-- Margaritifera_falcata **         
  |  |                                                
  |  |              +-- Lasmigona                     
  |  |           +--+                                 
  +--+           |  +-- Platynaias                    
     |  +--------+                                    
     |  |        |  +-- Pyganodon                     
     |  |        +--+                                 
     |  |           +-- Utterbackia                   
     +--+                                             
        |        +----- Fusconaia                     
        |  +-----+                                    
        |  |     |  +-- Pronodularia                  
        |  |     +--+                                 
        +--+        +-- Sinosolenaia                  
           |                                          
           |  +-------- Quadrula                      
           +--+                                       
              |  +----- Toxolasma                     
              +--+                                    
                 |  +-- Venustaconcha                 
                 +--+                                 
                    +-- Lampsilis

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

iTtTbiriSoTroTroSiD
2 2 0 2 90 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 6 0 0 -- 
  +-- Margaritifera_laevis **  
--+                         
  +-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
2 2 0 2 31 0 0 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 25 1 1 -- 
  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 25 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 55 0 0 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
2 2 0 2 90 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 51 0 0 -- 
  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 7 0 0 -- 
  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
2 2 0 2 35 0 0 -- 
  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
2 2 0 2 51 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 32 0 0 -- 
  +-- Margaritifera_middendorffi **
--+                             
  +-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 21 0 0 -- 
  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 10 0 0 -- 
  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 1 1 -- 
  +-- Pseudunio                  
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Margaritifera_laevis **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 53 0 0 -- 
-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 101 1 1 -- 
  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 1 1 -- 
  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 1 1 -- 
  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 2 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 0 0 -- 
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 0 0 -- 
-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 27 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 0 0 -- 
-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 39 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 2 2 -- 
  +----- Margaritifera_margaritifera **
--+                                 
  |  +-- Pseudunio                  
  +--+                              
     +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 79 1 1 -- 
  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 0 0 -- 
-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 1 1 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 2 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 8 0 0 -- 
-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 2 2 -- 
  +----- Pseudunio                  
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Cumberlandia

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 27 1 1 -- 
  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 0 0 -- 
-- Margaritifera_margaritifera **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 1 1 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 48 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 0 0 -- 
-- Margaritifera_falcata **

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 19 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 0 0 -- 
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
1 1 0 1 47 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 2 2 -- 
  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Pseudunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Margaritifera_dahurica **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 95 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 0 0 -- 
-- Margaritifera_dahurica **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 2 2 -- 
  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Pseudunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 0 0 -- 
-- Margaritifera_falcata **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 25 1 1 -- 
  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 1 1 -- 
  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 1 1 -- 
  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 0 0 -- 
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 0 0 -- 
-- Margaritifera_margaritifera **

Graf, 2000 [Publ 2647], fig. 1

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Margaritifera_dahurica **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 28 1 1 -- 
  +-- Cumberlandia         
--+                        
  +-- Margaritifera_falcata **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 4 0 0 -- 
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 2 2 -- 
  +----- Pseudunio                  
--+                                 
  |  +-- Cumberlandia               
  +--+                              
     +-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 50 0 0 -- 
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1 1 0 1 64 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 15 0 0 -- 
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 2 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 0 0 -- 
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 1 1 -- 
  +-- Pseudunio                  
--+                              
  +-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 101 1 1 -- 
  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia