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Species Richness & Endemism by
Geography

Data last updated
Mon 1 Dec 2025

Bivalvia | Unionoida | Unionoidea | Margaritiferidae | Margaritiferinae

genus Margaritifera Schumacher, 1816: cladomics

7 Recent valid species (23 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 (23 total)
analyzed in 220 cladograms in 128 publications
maximum number of terminals in an analysis = 7 (mean = 2.23, median = 2)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 111
maximum number of included terminals = 7, (mean = 3.44 median = 2)
proportion of cladograms that support basic monophyly = 0.95

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	61	3	2	11

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

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 **

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. 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 **

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. 4

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	61	3	2	11

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

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

tT	bi	riS	oT	roT	roS	iD
7	7	5	7	61	3	2	11

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

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., 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. 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. 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 **

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., 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 **

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 **

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. 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 **

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. 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 **

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 **

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 **

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 **

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. 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 **

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

tT	bi	riS	oT	roT	roS	iD
4	5	0	4	114	0	0	10

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

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 **

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 **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	71	2	2	9

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

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	93	2	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	93	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	71	2	2	9

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

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	71	2	2	9

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

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 **

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	85	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	34	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	83	2	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	35	1	1	9

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

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

tT	bi	riS	oT	roT	roS	iD
3	3	1	3	66	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 **

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 **

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

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

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

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	52	0	0	--

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

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 **

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

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

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

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., 2019 [Publ 3864], fig. 3

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

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

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

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

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

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 **

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 **

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

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

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

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 **

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 **

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 **

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 **

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 **

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 **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	36	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 **

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 **

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 **

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 **

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 **

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 **

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

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

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

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. 12

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

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

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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 **

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

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

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

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 **

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

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

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

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

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

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

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., 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., 2020 [Publ 4138], fig. 1

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

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

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 **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	74	0	0	--

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

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., 2020 [Publ 4121], fig. 3

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	47	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 **

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 **

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 **

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

tT	bi	riS	oT	roT	roS	iD
2	2	0	2	90	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 **

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 **

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

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

-- Margaritifera_margaritifera **

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., 2024 [Publ 4379], fig. 5

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

-- Margaritifera_margaritifera **

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

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

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

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

-- Margaritifera_hembeli **

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., 2018 [Publ 3758], fig. 5

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

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

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

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

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

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

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

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

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

-- Margaritifera_middendorffi **

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

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

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

-- Margaritifera_margaritifera **

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

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

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

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

-- 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

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

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

-- Margaritifera_hembeli **

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

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

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

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

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 **

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

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

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

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

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

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

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

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 **

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 **

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

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

-- Margaritifera_hembeli **

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., 2002 [Publ 2657], fig. 1

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_dahurica **

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	31	1	1	--

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

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_falcata **

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_falcata **

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 **

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

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

-- Margaritifera_margaritifera **

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

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

-- 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 **

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

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

-- Margaritifera_margaritifera **

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 **

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

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

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

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

-- Margaritifera_dahurica **

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

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

-- Margaritifera_dahurica **

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

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

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

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

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 **

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. 3A-M

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

-- Margaritifera_margaritifera **

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

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

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

-- Margaritifera_laevis **

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	45	1	1	--

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

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

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 **

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 **

Smith, 2021 [Publ 4331], fig. 1

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

-- Margaritifera_margaritifera **

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

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

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

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

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

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

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

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

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

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

-- Margaritifera_margaritifera **

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

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

-- Margaritifera_margaritifera **

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

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

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

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

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

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	13	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. 3-ML

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

-- 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 **

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

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

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

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

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

tT	bi	riS	oT	roT	roS	iD
1	1	0	1	51	0	0	--

-- Margaritifera_dahurica **

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

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

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

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

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

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

-- Margaritifera_margaritifera **

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

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

Graf, 2000 [Publ 2647], fig. 1

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

-- Margaritifera **

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

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

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

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

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 **

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

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