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 (29 total)
number of subtaxa (rank: genus) = 3
analyzed in 259 cladograms in 148 publications
maximum number of terminals in an analysis = 13 (mean = 2.8, median = 2)
maximum number of subtaxa (genus) = 3 (mean = 1.6, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 151
maximum number of included terminals = 13, (mean = 4.08 median = 2)
maximum number of subtaxa (genus) = 3 (mean = 2.03, median = 2)
proportion of cladograms that support basic monophyly = 0.87

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 41
(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 = 7.61, median = 9)
maximum number of subtaxa (genus) = 3 (mean = 2.9, median = 3)
proportion of cladograms that support rigorous monophyly = 0.71

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 **

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

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

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

iTtTbiriSoTroTroSiD
11 11 9 3 188 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. 2

iTtTbiriSoTroTroSiD
11 11 7 3 190 2 1 11 
  +-------- Gibbosula_rochechouartii
  |                                 
--+  +----- Gibbosula_laosensis     
  |  |                              
  +--+  +-- Cumberlandia **            
     |  |                           
     +--+-- Pseudunio **               
        |                           
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11 11 7 3 190 2 1 11 
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Cumberlandia **            
     +--+                              
        |  +-- Pseudunio **               
        +--+                           
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
10 10 8 3 19 2 1 14 
  +-------- 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 **

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

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

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

iTtTbiriSoTroTroSiD
10 10 8 3 3 1 1 12 
  +-------- 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 **

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. 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 **

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

iTtTbiriSoTroTroSiD
10 12 10 3 58 2 1 10 
  +-------- 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. S1

iTtTbiriSoTroTroSiD
9 10 8 3 130 1 1 12 
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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., 2017 [Publ 3761], fig. S2

iTtTbiriSoTroTroSiD
9 10 7 3 130 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 **

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

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

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

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

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

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. 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. 5-6

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

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. 1

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 **

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 **

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-ML

iTtTbiriSoTroTroSiD
5 5 3 3 69 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 **

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

iTtTbiriSoTroTroSiD
5 5 3 3 83 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 **

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

iTtTbiriSoTroTroSiD
5 5 3 3 81 1 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 **

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., 2022 [Publ 4238], fig. 3

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

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

iTtTbiriSoTroTroSiD
4 5 0 1 114 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 33 0 0 12 
  +-- Cumberlandia ** 
--+                
  +-- 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 34 0 0 14 
  +-- 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 **

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

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

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

iTtTbiriSoTroTroSiD
3 3 1 1 13 0 0 13 
1

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

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

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

iTtTbiriSoTroTroSiD
3 3 1 2 24 0 0 13 
  +-- 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

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 **

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 2 26 0 0 -- 
  +-- 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 **

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 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 **

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. 11

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 0 0 -- 
1

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

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

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 **

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 5 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 17 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 29 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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 51 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 31 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 19 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 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. 13

iTtTbiriSoTroTroSiD
2 2 0 1 8 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 90 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 7 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 14 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 90 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 30 0 0 -- 
  +-- Margaritifera **
--+                
  +-- Pseudunio **

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. 10

iTtTbiriSoTroTroSiD
2 2 0 1 12 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 74 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 55 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 9 0 0 -- 
1

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 6 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 4 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 25 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 120 0 0 -- 
1

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

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

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 **

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 37 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 35 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 1 21 0 0 -- 
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 51 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Bashê, 2022 [Publ 4265], fig. 1

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 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 47 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 103 0 0 -- 
-- Cumberlandia_monodonta **

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

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

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

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

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

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

Graf, 2002 [Publ 2651], fig. 2

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

Smith, 2021 [Publ 4331], fig. 1

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 48 1 1 -- 
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Margaritifera_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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Graf, 2000 [Publ 2647], fig. 1

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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. S1

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

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

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

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

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

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

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

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

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

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

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

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

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