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Page last updated
Thu 3 September 2025

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

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Bivalvia | Unionida | Unionoidea | Margaritiferidae | Margaritiferinae

genus Margaritifera Schumacher, 1815: cladomics
7 Recent valid species (26 total species)

Cladomics Summary

number of species = 7 (26 total)
analyzed in 231 cladograms in 135 publications
maximum number of terminals in an analysis = 7 (mean = 2.23, 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

NJ Tree

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

** Top Monophyly Cladogram **

tT	bi	riS	oT	roT	roS	iD
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 **

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	9	6	2	11

                 +-- Cumberlandia               
  +--------------+                              
  |              +-- Pseudunio                  
  |                                             
--+              +-- Margaritifera_margaritifera **
  |  +-----------+                              
  |  |           +-- Margaritifera_dahurica **     
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_laevis **       
        +--+                                    
           |  +----- Margaritifera_middendorffi ** 
           +--+                                 
              |  +-- Margaritifera_marrianae **    
              +--+                              
                 +-- Margaritifera_hembeli **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	27	3	2	11

              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_dahurica **     
  |  +--------+                              
  |  |        +-- Margaritifera_margaritifera **
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     |  |                                    
     +--+     +-- Margaritifera_hembeli **      
        |  +--+                              
        |  |  +-- Margaritifera_marrianae **    
        +--+                                 
           |  +-- Margaritifera_laevis **       
           +--+                              
              +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
7	7	3	7	194	4	2	10

  +-------------- Cumberlandia               
  |                                          
--+  +----------- Pseudunio                  
  |  |                                       
  |  |  +-------- Margaritifera_falcata **      
  +--+  |                                    
     |  |     +-- Margaritifera_dahurica **     
     |  +-----+                              
     +--+     +-- Margaritifera_margaritifera **
        |                                    
        |  +----- Margaritifera_laevis **       
        |  |                                 
        +--+----- Margaritifera_middendorffi ** 
           |                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	6	3	2	11

              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_dahurica **     
  |  +--------+                              
  |  |        +-- Margaritifera_margaritifera **
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     |  |                                    
     +--+     +-- Margaritifera_middendorffi ** 
        |  +--+                              
        |  |  +-- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_marrianae **    
           +--+                              
              +-- Margaritifera_hembeli **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	192	4	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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	21	4	2	11

              +-- Cumberlandia               
  +-----------+                              
  |           +-- Pseudunio                  
  |                                          
--+           +-- Margaritifera_dahurica **     
  |  +--------+                              
  |  |        +-- Margaritifera_margaritifera **
  +--+                                       
     |  +-------- Margaritifera_falcata **      
     |  |                                    
     +--+     +-- Margaritifera_middendorffi ** 
        |  +--+                              
        |  |  +-- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	19	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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	62	3	2	11

              +-- Margaritifera_margaritifera **
  +-----------+                              
  |           +-- Margaritifera_dahurica **     
--+                                          
  |  +----------- Margaritifera_falcata **      
  +--+                                       
     |  +-------- Margaritifera_middendorffi ** 
     +--+                                    
        |  +----- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	133	4	2	11

  +-------------------- Cumberlandia               
  |                                                
--+  +----------------- Pseudunio                  
  |  |                                             
  +--+              +-- Margaritifera_dahurica **     
     |  +-----------+                              
     |  |           +-- Margaritifera_margaritifera **
     +--+                                          
        |  +----------- Margaritifera_falcata **      
        +--+                                       
           |  +-------- Margaritifera_laevis **       
           +--+                                    
              |  +----- Margaritifera_middendorffi ** 
              +--+                                 
                 |  +-- Margaritifera_hembeli **      
                 +--+                              
                    +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	27	3	2	11

           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  |  |                                    
  +--+     +-- Margaritifera_hembeli **      
     |  +--+                              
     |  |  +-- Margaritifera_marrianae **    
     +--+                                 
        |  +-- Margaritifera_laevis **       
        +--+                              
           +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	62	3	2	11

              +-- Margaritifera_dahurica **     
  +-----------+                              
  |           +-- Margaritifera_margaritifera **
--+                                          
  |  +----------- Margaritifera_falcata **      
  +--+                                       
     |  +-------- Margaritifera_middendorffi ** 
     +--+                                    
        |  +----- Margaritifera_laevis **       
        +--+                                 
           |  +-- Margaritifera_hembeli **      
           +--+                              
              +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	4	3	2	11

           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  |  |                                    
  +--+     +-- Margaritifera_laevis **       
     |  +--+                              
     |  |  +-- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_marrianae **    
        +--+                              
           +-- Margaritifera_hembeli **

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

tT	bi	riS	oT	roT	roS	iD
7	7	4	7	194	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. 6-14

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	623	4	2	11

  +----------------- Cumberlandia               
  |                                             
--+              +-- Margaritifera_dahurica **     
  |  +-----------+                              
  |  |           +-- Margaritifera_margaritifera **
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_middendorffi ** 
        +--+                                    
           |  +----- Margaritifera_laevis **       
           +--+                                 
              |  +-- Margaritifera_hembeli **      
              +--+                              
                 +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	22	3	2	11

                 +-- Cumberlandia               
  +--------------+                              
  |              +-- Pseudunio                  
  |                                             
--+              +-- Margaritifera_margaritifera **
  |  +-----------+                              
  |  |           +-- Margaritifera_dahurica **     
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_laevis **       
        +--+                                    
           |  +----- Margaritifera_hembeli **      
           +--+                                 
              |  +-- Margaritifera_middendorffi ** 
              +--+                              
                 +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	62	3	2	11

           +-- Margaritifera_margaritifera **
  +--------+                              
  |        +-- Margaritifera_dahurica **     
--+                                       
  |  +-------- Margaritifera_falcata **      
  |  |                                    
  +--+     +-- Margaritifera_laevis **       
     |  +--+                              
     |  |  +-- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_hembeli **      
        +--+                              
           +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	21	4	2	11

                 +-- Cumberlandia               
  +--------------+                              
  |              +-- Pseudunio                  
  |                                             
--+              +-- Margaritifera_margaritifera **
  |  +-----------+                              
  |  |           +-- Margaritifera_dahurica **     
  +--+                                          
     |  +----------- Margaritifera_falcata **      
     +--+                                       
        |  +-------- Margaritifera_laevis **       
        +--+                                    
           |  +----- Margaritifera_middendorffi ** 
           +--+                                 
              |  +-- Margaritifera_hembeli **      
              +--+                              
                 +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
6	6	4	6	24	3	2	11

           +-- Margaritifera_margaritifera **
  +--------+                              
  |        +-- Margaritifera_dahurica **     
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_laevis **       
     +--+                                 
        |  +-- Margaritifera_marrianae **    
        +--+                              
           +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
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 **

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

tT	bi	riS	oT	roT	roS	iD
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

tT	bi	riS	oT	roT	roS	iD
6	6	4	6	133	3	2	11

           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_laevis **       
        +--+                              
           +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
6	6	4	6	70	3	2	11

           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_marrianae **    
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_laevis **

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

tT	bi	riS	oT	roT	roS	iD
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

tT	bi	riS	oT	roT	roS	iD
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 **

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

tT	bi	riS	oT	roT	roS	iD
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

tT	bi	riS	oT	roT	roS	iD
6	6	4	6	133	3	2	11

           +-- Margaritifera_dahurica **     
  +--------+                              
  |        +-- Margaritifera_margaritifera **
--+                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_middendorffi ** 
     +--+                                 
        |  +-- Margaritifera_laevis **       
        +--+                              
           +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
5	5	3	5	4	3	2	10

  +-------------- Cumberlandia               
  |                                          
--+  +----------- Pseudunio                  
  |  |                                       
  +--+        +-- Margaritifera_dahurica **     
     |  +-----+                              
     |  |     +-- Margaritifera_margaritifera **
     +--+                                    
        |  +----- Margaritifera_falcata **      
        +--+                                 
           |  +-- Margaritifera_middendorffi ** 
           +--+                              
              +-- Margaritifera_laevis **

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

tT	bi	riS	oT	roT	roS	iD
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. 4

tT	bi	riS	oT	roT	roS	iD
5	5	3	5	6	2	2	10

  +----------- Pseudunio                  
  |                                       
  +----------- Cumberlandia               
--+                                       
  |        +-- Margaritifera_dahurica **     
  |  +-----+                              
  |  |     +-- Margaritifera_margaritifera **
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
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

tT	bi	riS	oT	roT	roS	iD
5	5	3	5	4	3	2	10

  +-------------- Cumberlandia               
  |                                          
--+  +----------- Pseudunio                  
  |  |                                       
  +--+        +-- Margaritifera_dahurica **     
     |  +-----+                              
     |  |     +-- Margaritifera_margaritifera **
     +--+                                    
        |  +----- Margaritifera_falcata **      
        +--+                                 
           |  +-- Margaritifera_laevis **       
           +--+                              
              +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
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 **

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

tT	bi	riS	oT	roT	roS	iD
4	4	0	4	32	0	0	9

  +-- Margaritifera_dahurica **     
  |                              
  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_laevis **       
  |                              
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
4	4	2	4	32	2	2	9

  +----------- Cumberlandia               
  |                                       
--+  +-------- Pseudunio                  
  |  |                                    
  +--+     +-- Margaritifera_margaritifera **
     |  +--+                              
     |  |  +-- Margaritifera_dahurica **     
     +--+                                 
        |  +-- Margaritifera_falcata **      
        +--+                              
           +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
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

tT	bi	riS	oT	roT	roS	iD
4	5	0	4	113	0	0	10

  +-- Gibbosula                  
  |                              
  +-- Margaritifera_dahurica **     
  |                              
--+-- Margaritifera_margaritifera **
  |                              
  +-- Margaritifera_laevis **       
  |                              
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
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

tT	bi	riS	oT	roT	roS	iD
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. 4

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	65	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	84	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	35	1	1	9

  +-------- Cumberlandia               
--+                                    
  |  +----- Margaritifera_margaritifera **
  +--+                                 
     |  +-- Margaritifera_falcata **      
     +--+                              
        +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	82	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	70	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	92	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	70	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	7	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

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

  +----- Margaritifera_margaritifera **
--+                                 
  |  +-- Margaritifera_falcata **      
  +--+                              
     +-- Margaritifera_laevis **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	13	0	0	9

  +----- Margaritifera_margaritifera **
--+                                 
  |  +-- Margaritifera_falcata **      
  +--+                              
     +-- Margaritifera_marrianae **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	70	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	38	3	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
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-nt

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	92	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	38	3	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_dahurica **     
        +--+                              
           +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	39	2	2	9

  +----------- Cumberlandia               
--+                                       
  |  +-------- Pseudunio                  
  +--+                                    
     |  +----- Margaritifera_falcata **      
     +--+                                 
        |  +-- Margaritifera_margaritifera **
        +--+                              
           +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	36	1	1	9

  +-------- Cumberlandia               
--+                                    
  |  +----- Margaritifera_falcata **      
  +--+                                 
     |  +-- Margaritifera_marrianae **    
     +--+                              
        +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	36	1	1	--

  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	6	0	0	--

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	30	1	1	--

  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	4	19	0	0	--

  +-------- Margaritifera_laevis__2 **   
--+                                   
  |  +----- Margaritifera_laevis__1 **   
  +--+                                
     |  +-- Margaritifera_middendorffi **
     +--+                             
        +-- Margaritifera_laevis__3 **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	14	0	0	--

  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

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

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

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

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	12	0	0	--

  +-- Margaritifera_middendorffi **
--+                             
  +-- Margaritifera_laevis **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	17	0	0	--

  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_dahurica **

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

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

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	6	0	0	--

  +-- Margaritifera_laevis **  
--+                         
  +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	12	0	0	--

  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_falcata **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_hembeli **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	34	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	9	0	0	--

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_hembeli **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	8	0	0	--

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	25	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	26	1	1	--

  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	34	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	34	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	6	0	0	--

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	51	0	0	--

  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	7	0	0	--

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	38	1	1	--

  +----- Cumberlandia          
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

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

  +----- Cumberlandia          
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	17	1	1	--

  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	42	1	1	--

  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_dahurica **
  +--+                         
     +-- Margaritifera_falcata **

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

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

  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
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

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	3	12	1	1	--

  +----- Cumberlandia               
  |                                 
--+  +-- Margaritifera_margaritifera **
  |  |                              
  +--+-- Margaritifera_dahurica__1 **  
     |                              
     +-- Margaritifera_dahurica__2 **

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

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

  +----- Cumberlandia          
--+                            
  |  +-- Margaritifera_dahurica **
  +--+                         
     +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	10	0	0	--

  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	34	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	21	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	42	1	1	--

  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_dahurica **
  +--+                         
     +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	32	0	0	--

  +-- Margaritifera_middendorffi **
--+                             
  +-- Margaritifera_laevis **

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

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

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	10	0	0	--

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	12	0	0	--

  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

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

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	34	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	10	0	0	--

  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	25	1	1	--

  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	35	0	0	--

  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	34	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_falcata ** 
--+                         
  +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	37	0	0	--

  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_falcata **

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

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

  +-- Margaritifera_falcata **      
--+                              
  +-- Margaritifera_margaritifera **

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

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

  +----- Pseudunio             
--+                            
  |  +-- Margaritifera_falcata ** 
  +--+                         
     +-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	73	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_dahurica **
--+                         
  +-- Margaritifera_laevis **

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

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

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

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

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	31	0	0	--

  +-- Margaritifera_margaritifera **
--+                              
  +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	51	0	0	--

  +-- Margaritifera_dahurica **     
--+                              
  +-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	31	0	0	--

  +-- Margaritifera_laevis **      
--+                             
  +-- Margaritifera_middendorffi **

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

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

-- Margaritifera_laevis **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	48	0	0	--

-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	27	0	0	--

-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

  +-- Cumberlandia         
--+                        
  +-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	39	1	1	--

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

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

-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	19	0	0	--

-- Margaritifera_falcata **

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

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

-- Margaritifera_margaritifera **

Smith, 2021 [Publ 4331], fig. 1

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	19	0	0	--

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	6	0	0	--

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	27	1	1	--

  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	43	0	0	--

-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	80	1	1	--

  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	52	0	0	--

-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	15	0	0	--

-- Margaritifera_falcata **

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

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

-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	21	0	0	--

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	21	2	2	--

  +----- Pseudunio                  
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Cumberlandia

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

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

  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	19	0	0	--

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	21	2	2	--

  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Pseudunio

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

  +----- Margaritifera_margaritifera **
--+                                 
  |  +-- Pseudunio                  
  +--+                              
     +-- Cumberlandia

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	35	1	1	--

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

-- Margaritifera_dahurica **

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

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

  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_dahurica **

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

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

  +-- Pseudunio                  
--+                              
  +-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

Graf, 2000 [Publ 2647], fig. 1

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

-- Margaritifera **

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	11	0	0	--

-- Margaritifera_hembeli **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	63	0	0	--

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	19	0	0	--

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	52	0	0	--

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	18	1	1	--

  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

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

-- Margaritifera_dahurica **

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

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

  +----- Cumberlandia               
--+                                 
  |  +-- Margaritifera_margaritifera **
  +--+                              
     +-- Pseudunio

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	21	0	0	--

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

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

-- Margaritifera_dahurica **

Wu et al., 2026 [Publ 4825], fig. 6B

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	64	0	0	--

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	50	0	0	--

-- Margaritifera_dahurica **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	25	1	1	--

  +-- Margaritifera_falcata **
--+                        
  +-- Cumberlandia

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	6	0	0	--

-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	35	1	1	--

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

-- Margaritifera_hembeli **

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

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

-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

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

  +-- Pseudunio                  
--+                              
  +-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	101	1	1	--

  +-- Margaritifera_margaritifera **
--+                              
  +-- Cumberlandia

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

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

-- Margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	6	0	0	--

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_falcata **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	52	0	0	--

-- Margaritifera_dahurica **

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

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

  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

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

  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	6	0	0	--

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

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

  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

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

  +----- Pseudunio                  
--+                                 
  |  +-- Cumberlandia               
  +--+                              
     +-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_hembeli **

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

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

-- Margaritifera_middendorffi **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	101	1	1	--

  +-- Cumberlandia               
--+                              
  +-- Margaritifera_margaritifera **

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

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

  +-- Margaritifera_margaritifera **
--+                              
  +-- Pseudunio

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	19	0	0	--

-- Margaritifera_margaritifera **

Kashiri et al., 2024 [Publ 4822], fig. 3

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	43	0	0	--

-- Margaritifera_dahurica **

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