Bivalvia | Unionida | Unionoidea | Margaritiferidae

subfamily MARGARITIFERINAE Henderson, 1929: cladomics

3 genera

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 = 11 (32 total)
number of subtaxa (rank: genus) = 3
analyzed in 264 cladograms in 150 publications
maximum number of terminals in an analysis = 13 (mean = 2.84, median = 2)
maximum number of subtaxa (genus) = 3 (mean = 1.61, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 154
maximum number of included terminals = 13, (mean = 4.16 median = 2)
maximum number of subtaxa (genus) = 3 (mean = 2.05, median = 2)
proportion of cladograms that support basic monophyly = 0.87

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 43
(Rigorous monophyly refers to cladograms with terminal species representing more than one subtaxon (genus) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 13 (mean = 7.77, median = 9)
maximum number of subtaxa (genus) = 3 (mean = 2.91, median = 3)
proportion of cladograms that support rigorous monophyly = 0.7

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 Margaritiferinae cladogram distances
PCoA Ordination

NJ Tree of Margaritiferinae cladogram distances
NJ Tree

Cladograms

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

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

iTtTbiriSoTroTroSiD
11 15 13 3 15 4 1 0 
  +----------- Pseudunio_marocanus **      
  |                                     
  |        +-- Pseudunio_auricularius__1 **
--+        |                            
  |  +-----+-- Pseudunio_auricularius__2 **
  |  |     |                            
  +--+     +-- Pseudunio_homsensis **      
     |                                  
     |  +----- Gibbosulinae             
     +--+                               
        |  +-- Cumberlandia **             
        +--+                            
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
13 13 9 3 3 3 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
11 11 9 3 130 3 1 11 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11 11 7 3 189 2 1 11 
  +-------- Gibbosula_rochechouartii
  |                                 
--+  +----- Gibbosula_laosensis     
  |  |                              
  +--+  +-- Cumberlandia **            
     |  |                           
     +--+-- Pseudunio **               
        |                           
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11 14 12 3 620 3 1 4 
  +-------- Pseudunio **    
--+                      
  |  +----- Gibbosulinae 
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11 11 9 3 187 2 1 11 
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Margaritifera **           
     +--+                              
        |  +-- Cumberlandia **            
        +--+                           
           +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
11 11 9 3 17 3 1 11 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
11 11 7 3 189 2 1 11 
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Cumberlandia **            
     +--+                              
        |  +-- Pseudunio **               
        +--+                           
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11 11 9 3 17 3 1 11 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10 12 10 3 58 2 1 10 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
10 12 10 3 24 2 1 12 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10 10 8 3 24 2 1 12 
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Margaritifera **           
     +--+                              
        |  +-- Cumberlandia **            
        +--+                           
           +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10 12 10 3 58 2 1 10 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10 10 8 3 3 1 1 12 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10 10 8 3 19 2 1 14 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10 12 10 3 58 2 1 12 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
10 10 8 3 3 1 1 12 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10 11 9 3 1 1 1 13 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
9 10 8 3 67 1 1 14 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
9 10 7 3 129 1 1 11 
  +----- Cumberlandia ** 
  |                   
--+  +-- Gibbosulinae 
  |  |                
  +--+-- Pseudunio **    
     |                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
9 10 8 3 21 1 1 12 
  +-------- Pseudunio **    
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
9 10 8 3 67 1 1 12 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Gibbosulinae 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
9 10 8 3 21 1 1 12 
  +-------- Pseudunio **    
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
9 10 8 3 129 1 1 12 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
8 8 5 3 1 0 0 16 
  +----------- Margaritifera_margaritifera **
  |                                       
  |     +----- Margaritifera_dahurica__1 **  
  |     |                                 
  +-----+----- Margaritifera_dahurica__2 **  
--+     |                                 
  |     |  +-- Pseudunio **                  
  |     +--+                              
  |        +-- Cumberlandia **               
  |                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_laevis **       
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
8 8 6 3 1 0 0 12 
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
8 8 6 3 1 0 0 12 
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
8 8 6 3 2 0 0 8 
  +-------- Pseudunio_marocanus **   
--+                               
  |  +----- Pseudunio_auricularius **
  +--+                            
     |  +-- Cumberlandia **          
     +--+                         
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
8 8 6 3 2 0 0 10 
  +----- Pseudunio_auricularius__1 **
  |                               
--+----- Pseudunio_auricularius__2 **
  |                               
  |  +-- Cumberlandia **             
  +--+                            
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
7 7 4 3 4 0 0 10 
  +-- Pseudunio **    
  |                
--+-- Cumberlandia ** 
  |                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
7 7 5 3 5 0 0 11 
  +----- Margaritifera **
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
7 7 5 3 1 0 0 13 
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
7 7 3 3 5 0 0 14 
  +-------- Margaritifera_dahurica **        
  |                                       
  +-------- Margaritifera_margaritifera__1 **
  |                                       
  +-------- Margaritifera_margaritifera__2 **
  |                                       
--+-------- Margaritifera_margaritifera__3 **
  |                                       
  |     +-- Pseudunio **                     
  +-----+                                 
  |     +-- Cumberlandia **                  
  |                                       
  |  +----- Margaritifera_falcata **         
  +--+                                    
     |  +-- Margaritifera_middendorffi **    
     +--+                                 
        +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6 6 4 3 30 0 0 12 
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
6 6 4 3 35 1 1 14 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
6 6 4 3 85 1 1 12 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
6 6 2 3 2 0 0 18 
  +-------- Margaritifera_margaritifera **  
  |                                      
  |  +----- Cumberlandia **                 
--+  |                                   
  |  |  +-- Pseudunio **                    
  |  |  |                                
  +--+  +-- Margaritifera_dahurica **       
     |  |                                
     |  +-- Margaritifera_middendorffi__1 **
     |  |                                
     |  +-- Margaritifera_middendorffi__2 **
     +--+                                
        +-- Margaritifera_falcata__1 **     
        |                                
        +-- Margaritifera_falcata__2 **     
        |                                
        +-- Margaritifera_falcata__3 **     
        |                                
        +-- Margaritifera_falcata__4 **

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

iTtTbiriSoTroTroSiD
6 6 4 3 35 1 1 14 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
6 6 4 3 85 1 1 12 
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
5 5 3 3 91 2 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 69 2 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 69 1 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 51 1 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 91 2 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 37 1 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 69 2 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 5 2 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 64 2 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 83 1 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5 5 3 3 81 1 1 13 
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
4 4 0 1 32 0 0 14 
1

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

iTtTbiriSoTroTroSiD
4 4 2 2 35 0 0 14 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
4 5 0 1 113 1 1 14 
  +-- Gibbosulinae               
  |                              
  +-- Margaritifera_dahurica **     
  |                              
--+-- Margaritifera_margaritifera **
  |                              
  +-- Margaritifera_laevis **       
  |                              
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
4 4 2 2 34 0 0 12 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 19 5 2 16 0 0 28 
  +----------- Cumberlandia **       
  |                               
  +----------- Margaritifera **      
  |                               
  |  +-------- Gonideinae         
--+  |                            
  |  |     +-- Elliptio           
  |  |     |                      
  |  |     +-- Fusconaia          
  |  |     |                      
  +--+  +--+-- Amblemini          
     |  |  |                      
     |  |  +-- Pleurobema         
     |  |  |                      
     |  |  +-- Unionini           
     |  |                         
     |  |  +-- Pyganodon_cataracta
     +--+  |                      
        |  +-- Pyganodon_grandis  
        |  |                      
        |  +-- Utterbackia        
        |  |                      
        |  +-- Lampsilis_teres    
        |  |                      
        |  +-- Lampsilis_straminea
        +--+                      
           +-- Megalonaias        
           |                      
           +-- Obliquaria         
           |                      
           +-- Theliderma         
           |                      
           +-- Quadrula           
           |                      
           +-- Uniomerus

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

iTtTbiriSoTroTroSiD
3 3 1 2 41 0 0 13 
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 19 0 0 15 
  +----- Cumberlandia ** 
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
3 3 1 3 3 0 0 15 
  +----- Margaritifera **
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
3 3 1 1 0 0 0 13 
1

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

iTtTbiriSoTroTroSiD
3 3 1 2 43 1 1 13 
  +----- Gibbosulinae 
--+                   
  |  +-- Cumberlandia ** 
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 2 41 0 0 13 
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 1 0 0 15 
  +----- Cumberlandia ** 
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
3 3 1 2 46 1 1 13 
  +----- Gibbosulinae 
--+                   
  |  +-- Cumberlandia ** 
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 2 24 0 0 13 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 2 35 0 0 13 
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 22 4 2 19 0 0 30 
  +----------- Cumberlandia **       
  |                               
  +----------- None__2            
  |                               
  +----------- Margaritifera **      
--+                               
  |  +-------- None__1            
  |  |                            
  |  |  +----- Anodontina         
  |  |  |                         
  |  |  |  +-- Gonideinae         
  +--+  |  |                      
     |  |  +-- Pyganodon_grandis  
     |  |  |                      
     |  |  +-- Utterbackia        
     |  |  |                      
     |  |  +-- Pyganodon_cataracta
     +--+  |                      
        |  +-- Lampsilis_straminea
        |  |                      
        |  +-- Lampsilis_teres    
        |  |                      
        |  +-- Megalonaias        
        |  |                      
        |  +-- Obliquaria         
        |  |                      
        +--+-- Theliderma         
           |                      
           +-- Quadrula           
           |                      
           +-- Uniomerus          
           |                      
           +-- Amblema            
           |                      
           +-- Elliptio           
           |                      
           +-- Fusconaia          
           |                      
           +-- Plectomerus        
           |                      
           +-- Pleurobema         
           |                      
           +-- Unionini

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

iTtTbiriSoTroTroSiD
3 3 1 2 37 0 0 13 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 19 0 0 13 
  +----- Pseudunio **    
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
3 3 1 2 29 0 0 13 
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 3 44 0 0 13 
  +----- Pseudunio **    
--+                   
  |  +-- Cumberlandia ** 
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 22 3 2 19 0 0 30 
  +-------- Cumberlandia **       
  |                            
  +-------- None__1            
  |                            
  +-------- None__2            
  |                            
  +-------- Margaritifera **      
  |                            
--+  +----- Anodontina         
  |  |                         
  |  |  +-- Gonideinae         
  |  |  |                      
  |  |  +-- Pyganodon_grandis  
  |  |  |                      
  |  |  +-- Utterbackia        
  |  |  |                      
  |  |  +-- Pyganodon_cataracta
  +--+  |                      
     |  +-- Lampsilis_straminea
     |  |                      
     |  +-- Lampsilis_teres    
     |  |                      
     |  +-- Megalonaias        
     |  |                      
     |  +-- Obliquaria         
     |  |                      
     +--+-- Theliderma         
        |                      
        +-- Quadrula           
        |                      
        +-- Uniomerus          
        |                      
        +-- Amblema            
        |                      
        +-- Elliptio           
        |                      
        +-- Fusconaia          
        |                      
        +-- Plectomerus        
        |                      
        +-- Aculamprotulina    
        |                      
        +-- Unionina

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

iTtTbiriSoTroTroSiD
3 3 1 2 11 0 0 14 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 1 13 0 0 13 
1

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

iTtTbiriSoTroTroSiD
3 3 1 2 45 0 0 13 
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3 3 1 2 25 0 0 13 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
2 2 0 1 17 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 51 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 28 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 25 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 90 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 34 0 2 32 0 0 45 
  +----------------- Cumberlandia **   
  |                                 
--+----------------- Margaritifera **  
  |                                 
  |  +-------------- Unioninae      
  +--+                              
     |  +----------- Sinosolenaia   
     +--+                           
        |  +-------- Lamprotulini   
        +--+                        
           |  +----- Chamberlainiini
           +--+                     
              |  +-- Ptychorhynchus 
              +--+                  
                 +-- Ambleminae

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

iTtTbiriSoTroTroSiD
2 2 0 1 7 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 19 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 17 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 37 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 24 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 18 0 0 -- 
  +-- Pseudunio **   
--+               
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 120 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 52 1 1 -- 
  +----- Gibbosulinae
--+                  
  |  +-- Pseudunio **   
  +--+               
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 45 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 12 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 100 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 1 12 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 43 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Pseudunio **

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 2 23 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 1 5 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 38 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 44 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 1 74 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 90 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 55 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 26 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 31 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 34 0 2 32 0 0 45 
  +-------- Cumberlandia **   
  |                        
  +-------- Margaritifera **  
--+                        
  |  +----- Unioninae      
  |  |                     
  |  |  +-- Ptychorhynchus 
  +--+  |                  
     |  +-- Chamberlainiini
     |  |                  
     +--+-- Lamprotulini   
        |                  
        +-- Sinosolenaia   
        |                  
        +-- Ambleminae

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

iTtTbiriSoTroTroSiD
2 2 0 1 51 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 45 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 2 78 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 1 32 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 31 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 17 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 27 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 1 14 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 19 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 21 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 15 0 0 -- 
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 100 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 12 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 1 8 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 23 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 2 28 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 15 0 1 14 0 0 27 
  +-------------- Margaritifera_margaritifera__2 **
  |                                             
--+           +-- Margaritifera_margaritifera__1 **
  |  +--------+                                 
  |  |        +-- Margaritifera_falcata **         
  +--+                                          
     |  +-------- Unioninae                     
     |  |                                       
     +--+     +-- Pleurobemini                  
        |  +--+                                 
        |  |  +-- Gonideinae                    
        +--+                                    
           |  +-- Lampsilini                    
           +--+                                 
              +-- Quadrulini

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 21 0 0 -- 
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2 2 0 2 44 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2 2 0 1 55 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 9 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 32 0 0 -- 
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 56 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 0 0 -- 
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

Bashê, 2022 [Publ 4265], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 5 0 0 -- 
-- Pseudunio_homsensis **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 104 0 0 -- 
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Cumberlandia_monodonta **

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 17 0 0 -- 
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 0 0 -- 
-- Pseudunio_auricularius **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 94 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 0 0 -- 
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Cumberlandia_monodonta **

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

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

Smith, 2021 [Publ 4331], fig. 1

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 47 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Cumberlandia_monodonta **

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 55 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 0 0 -- 
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 0 0 -- 
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 0 0 -- 
-- Pseudunio_marocanus **

Graf, 2000 [Publ 2647], fig. 1

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 0 0 -- 
-- Pseudunio_marocanus **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 53 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 50 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 0 0 -- 
-- Cumberlandia_monodonta **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1 1 0 1 64 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 0 0 -- 
-- Pseudunio_auricularius **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 0 0 -- 
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 0 0 -- 
-- Cumberlandia_monodonta **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 17 0 0 -- 
-- Pseudunio_auricularius **