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 266 cladograms in 151 publications
maximum number of terminals in an analysis = 13 (mean = 2.83, 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. 6-14

iTtTbiriSoTroTroSiD
11 14 12 3 621 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 **

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. 1

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

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. 3_5

iTtTbiriSoTroTroSiD
11 11 9 3 130 3 1 11 
  +-------- Gibbosulinae 
--+                      
  |  +----- 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 **

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

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 **

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 **

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

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

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

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

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

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

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

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., 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. 4

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 **

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

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

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 **

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 **

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. 5-6

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

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 **

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. 4

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

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

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

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

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

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

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

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

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

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 **

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 **

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

iTtTbiriSoTroTroSiD
5 5 3 3 91 2 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 **

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-aa

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-BI

iTtTbiriSoTroTroSiD
5 5 3 3 69 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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
3 3 1 1 13 0 0 13 
1

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

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

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 **

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

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

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

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

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

iTtTbiriSoTroTroSiD
3 3 1 2 46 1 1 13 
  +----- Gibbosulinae 
--+                   
  |  +-- 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

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

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

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

iTtTbiriSoTroTroSiD
3 3 1 2 11 0 0 14 
  +-- 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

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

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

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

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

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

iTtTbiriSoTroTroSiD
3 3 1 2 35 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 **

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 2 21 0 0 -- 
  +-- Pseudunio **    
--+                
  +-- 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. 6

iTtTbiriSoTroTroSiD
2 2 0 1 9 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 14 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 **

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

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

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

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

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

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

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

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

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

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. 2-4

iTtTbiriSoTroTroSiD
2 2 0 1 120 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 74 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

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

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

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

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. 16

iTtTbiriSoTroTroSiD
2 2 0 1 6 0 0 -- 
1

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 19 0 0 -- 
1

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 37 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 31 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 17 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 5 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 29 0 0 -- 
  +-- 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

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 8 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 34 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. 8a_S1

iTtTbiriSoTroTroSiD
2 2 0 1 90 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 31 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 25 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 55 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 90 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 51 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 7 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 51 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 34 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 21 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Bashê, 2022 [Publ 4265], fig. 1

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Klymus et al., 2025 [Publ 4814], fig. 3B

iTtTbiriSoTroTroSiD
1 1 0 1 35 0 0 -- 
-- Pseudunio_marocanus **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Smith, 2021 [Publ 4331], fig. 1

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 95 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Margaritifera_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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Graf, 2000 [Publ 2647], fig. 1

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

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

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

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

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

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. 7A

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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 **

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

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

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. 4

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

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

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

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

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

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

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

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

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

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

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

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

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

Klymus et al., 2025 [Publ 4814], fig. 3A

iTtTbiriSoTroTroSiD
1 1 0 1 35 0 0 -- 
-- Pseudunio_marocanus **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 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 **