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Page last updated
Thu 3 September 2025

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

FM(U)otW(aolcb) is the web version of the MUSSEL Project Database. Follow the links to browse the data or use the custom Google search field. Either way, you win!

Search the database pages:

Bivalvia | Unionida | Unionoidea | Margaritiferidae

subfamily MARGARITIFERINAE Henderson, 1929: cladomics
     3 genera

Cladomics Summary

number of species = 11 (32 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


NJ Tree

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

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

iTtTbiriSoTroTroSiD
111513315410
  +----------- Pseudunio_marocanus **      
  |                                     
  |        +-- Pseudunio_auricularius__1 **
--+        |                            
  |  +-----+-- Pseudunio_auricularius__2 **
  |  |     |                            
  +--+     +-- Pseudunio_homsensis **      
     |                                  
     |  +----- Gibbosulinae             
     +--+                               
        |  +-- Cumberlandia **             
        +--+                            
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
13139333113
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
111193173111
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1111731902111
  +-------- Gibbosula_rochechouartii
  |                                 
--+  +----- Gibbosula_laosensis     
  |  |                              
  +--+  +-- Cumberlandia **            
     |  |                           
     +--+-- Pseudunio **               
        |                           
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1111931882111
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Margaritifera **           
     +--+                              
        |  +-- Cumberlandia **            
        +--+                           
           +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
111193173111
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1111731902111
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Cumberlandia **            
     +--+                              
        |  +-- Pseudunio **               
        +--+                           
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
101083242112
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Margaritifera **           
     +--+                              
        |  +-- Cumberlandia **            
        +--+                           
           +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1012103582112
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
10108331112
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10119311113
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1012103582110
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1012103242112
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1012103582110
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
101083192114
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10108331112
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
91083671112
  +-------- Cumberlandia ** 
--+                      
  |  +----- Gibbosulinae 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
91083211112
  +-------- Pseudunio **    
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
910731301111
  +----- Cumberlandia ** 
  |                   
--+  +-- Gibbosulinae 
  |  |                
  +--+-- Pseudunio **    
     |                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
91083211112
  +-------- Pseudunio **    
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
91083671114
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
910831301112
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
885310016
  +----------- 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
886310012
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
886310012
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
886320010
  +----- Pseudunio_auricularius__1 **
  |                               
--+----- Pseudunio_auricularius__2 **
  |                               
  |  +-- Cumberlandia **             
  +--+                            
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
88632008
  +-------- Pseudunio_marocanus **   
--+                               
  |  +----- Pseudunio_auricularius **
  +--+                            
     |  +-- Cumberlandia **          
     +--+                         
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
775350011
  +----- Margaritifera **
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
774340010
  +-- Pseudunio **    
  |                
--+-- Cumberlandia ** 
  |                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
775310013
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
773350014
  +-------- Margaritifera_dahurica **        
  |                                       
  +-------- Margaritifera_margaritifera__1 **
  |                                       
  +-------- Margaritifera_margaritifera__2 **
  |                                       
--+-------- Margaritifera_margaritifera__3 **
  |                                       
  |     +-- Pseudunio **                     
  +-----+                                 
  |     +-- Cumberlandia **                  
  |                                       
  |  +----- Margaritifera_falcata **         
  +--+                                    
     |  +-- Margaritifera_middendorffi **    
     +--+                                 
        +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
6643351114
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
6643851112
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
6643851112
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
6643351114
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
6643300012
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
662320018
  +-------- Margaritifera_margaritifera **  
  |                                      
  |  +----- Cumberlandia **                 
--+  |                                   
  |  |  +-- Pseudunio **                    
  |  |  |                                
  +--+  +-- Margaritifera_dahurica **       
     |  |                                
     |  +-- Margaritifera_middendorffi__1 **
     |  |                                
     |  +-- Margaritifera_middendorffi__2 **
     +--+                                
        +-- Margaritifera_falcata__1 **     
        |                                
        +-- Margaritifera_falcata__2 **     
        |                                
        +-- Margaritifera_falcata__3 **     
        |                                
        +-- Margaritifera_falcata__4 **

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

iTtTbiriSoTroTroSiD
553352113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533912113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533831113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533811113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533642113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533691113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533371113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533511113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533692113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533912113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
5533692113
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
4422340014
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
4401320014
1

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

iTtTbiriSoTroTroSiD
4422330012
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
45011141114
  +-- Gibbosulinae               
  |                              
  +-- Margaritifera_dahurica **     
  |                              
--+-- Margaritifera_margaritifera **
  |                              
  +-- Margaritifera_laevis **       
  |                              
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
32232190030
  +-------- 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

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

iTtTbiriSoTroTroSiD
3311130013
1

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

iTtTbiriSoTroTroSiD
3312410013
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331100013
1

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

iTtTbiriSoTroTroSiD
3312431113
  +----- Gibbosulinae 
--+                   
  |  +-- Cumberlandia ** 
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3312461113
  +----- Gibbosulinae 
--+                   
  |  +-- Cumberlandia ** 
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
31952160028
  +----------- Cumberlandia **       
  |                               
  +----------- Margaritifera **      
  |                               
  |  +-------- Gonideinae         
--+  |                            
  |  |     +-- Elliptio           
  |  |     |                      
  |  |     +-- Fusconaia          
  |  |     |                      
  +--+  +--+-- Amblemini          
     |  |  |                      
     |  |  +-- Pleurobema         
     |  |  |                      
     |  |  +-- Unionini           
     |  |                         
     |  |  +-- Pyganodon_cataracta
     +--+  |                      
        |  +-- Pyganodon_grandis  
        |  |                      
        |  +-- Utterbackia        
        |  |                      
        |  +-- Lampsilis_teres    
        |  |                      
        |  +-- Lampsilis_straminea
        +--+                      
           +-- Megalonaias        
           |                      
           +-- Obliquaria         
           |                      
           +-- Theliderma         
           |                      
           +-- Quadrula           
           |                      
           +-- Uniomerus

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

iTtTbiriSoTroTroSiD
3312250013
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3313190015
  +----- Cumberlandia ** 
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
3312450013
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331330015
  +----- Margaritifera **
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
3312110014
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3312350013
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331310015
  +----- Cumberlandia ** 
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
3312370013
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3312240013
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3313190013
  +----- Pseudunio **    
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
3312410013
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3312290013
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
32242190030
  +----------- 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

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

iTtTbiriSoTroTroSiD
23402320045
  +-------- Cumberlandia **   
  |                        
  +-------- Margaritifera **  
--+                        
  |  +----- Unioninae      
  |  |                     
  |  |  +-- Ptychorhynchus 
  +--+  |                  
     |  +-- Chamberlainiini
     |  |                  
     +--+-- Lamprotulini   
        |                  
        +-- Sinosolenaia   
        |                  
        +-- Ambleminae

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

iTtTbiriSoTroTroSiD
22024500--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22022300--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22013100--
1

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

iTtTbiriSoTroTroSiD
220210000--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22022900--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22023000--
  +-- Margaritifera **
--+                
  +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
22013100--
1

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
22022100--
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22024400--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22022700--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22011400--
1

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

iTtTbiriSoTroTroSiD
22023200--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22021700--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22022300--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22017400--
1

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

iTtTbiriSoTroTroSiD
22012100--
1

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
2202100--
  +-- Margaritifera **
--+                
  +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
22022900--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22023400--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22019000--
1

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

iTtTbiriSoTroTroSiD
21501140027
  +-------------- Margaritifera_margaritifera__2 **
  |                                             
--+           +-- Margaritifera_margaritifera__1 **
  |  +--------+                                 
  |  |        +-- Margaritifera_falcata **         
  +--+                                          
     |  +-------- Unioninae                     
     |  |                                       
     +--+     +-- Pleurobemini                  
        |  +--+                                 
        |  |  +-- Gonideinae                    
        +--+                                    
           |  +-- Lampsilini                    
           +--+                                 
              +-- Quadrulini

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

iTtTbiriSoTroTroSiD
22022600--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2201600--
1

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

iTtTbiriSoTroTroSiD
22011200--
1

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

iTtTbiriSoTroTroSiD
22022900--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22022800--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2201800--
1

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

iTtTbiriSoTroTroSiD
22021500--
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
22011700--
1

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

iTtTbiriSoTroTroSiD
22019000--
1

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

iTtTbiriSoTroTroSiD
23402320045
  +----------------- Cumberlandia **   
  |                                 
--+----------------- Margaritifera **  
  |                                 
  |  +-------------- Unioninae      
  +--+                              
     |  +----------- Sinosolenaia   
     +--+                           
        |  +-------- Lamprotulini   
        +--+                        
           |  +----- Chamberlainiini
           +--+                     
              |  +-- Ptychorhynchus 
              +--+                  
                 +-- Ambleminae

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

iTtTbiriSoTroTroSiD
22015100--
1

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

iTtTbiriSoTroTroSiD
22011000--
1

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

iTtTbiriSoTroTroSiD
2201600--
1

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
22012900--
1

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

iTtTbiriSoTroTroSiD
22011900--
1

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

iTtTbiriSoTroTroSiD
22021800--
  +-- Pseudunio **   
--+               
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22015500--
1

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

iTtTbiriSoTroTroSiD
22011211--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22012900--
1

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

iTtTbiriSoTroTroSiD
22024400--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22015100--
1

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

iTtTbiriSoTroTroSiD
22022800--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22022900--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22013700--
1

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

iTtTbiriSoTroTroSiD
2201400--
1

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

iTtTbiriSoTroTroSiD
22023400--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22011700--
1

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

iTtTbiriSoTroTroSiD
22025211--
  +----- Gibbosulinae
--+                  
  |  +-- Pseudunio **   
  +--+               
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
2201400--
1

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
22012500--
1

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

iTtTbiriSoTroTroSiD
2201400--
1

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

iTtTbiriSoTroTroSiD
22022900--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22027800--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
22011200--
1

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

iTtTbiriSoTroTroSiD
220210000--
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
2201700--
1

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

iTtTbiriSoTroTroSiD
22024500--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22015500--
1

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

iTtTbiriSoTroTroSiD
2201900--
1

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

iTtTbiriSoTroTroSiD
22024400--
  +-- Margaritifera **
--+                
  +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
2202100--
  +-- Margaritifera **
--+                
  +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
22011000--
1

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

iTtTbiriSoTroTroSiD
2201600--
1

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

iTtTbiriSoTroTroSiD
2201500--
1

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

iTtTbiriSoTroTroSiD
22023800--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22022400--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22013211--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
220112000--
1

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

iTtTbiriSoTroTroSiD
22011000--
1

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

iTtTbiriSoTroTroSiD
22021911--
  +-- Margaritifera **
--+                
  +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

Graf, 2000 [Publ 2647], fig. 1

iTtTbiriSoTroTroSiD
11011600--
-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
110110300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11013011--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011600--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11014411--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
11012700--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011400--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011800--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
11011000--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1101700--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011700--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11011000--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
11014711--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11012111--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1101900--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
11011600--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11014411--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
11011500--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
11011400--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101800--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11014611--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11015600--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11016411--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101711--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11014611--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
11011400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
11011700--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

Bashê, 2022 [Publ 4265], fig. 1

iTtTbiriSoTroTroSiD
1101500--
-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1101500--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101700--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11014811--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11012111--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1101900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11014411--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1101911--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1101300--
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11012200--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
11014411--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11013311--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101200--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11015500--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11012900--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1101900--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
1101900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101911--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11013011--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1101300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11014411--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11014811--
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011100--
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
11011300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011800--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
11011900--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011300--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
11011000--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101700--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11013011--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101900--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011400--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11015311--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11015111--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
1101900--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
11011000--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11013000--
-- Margaritifera_margaritifera **
 
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