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Page last updated
Thu 17 July 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!

Bivalvia | Unionoida | Unionoidea | Margaritiferidae

subfamily MARGARITIFERINAE Henderson, 1929: cladomics
     3 genera

Cladomics Summary

number of species = 11 (29 total)
number of subtaxa (rank: genus) = 3
analyzed in 251 cladograms in 141 publications
maximum number of terminals in an analysis = 13 (mean = 2.79, 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) = 148
maximum number of included terminals = 13, (mean = 4.03 median = 2)
maximum number of subtaxa (genus) = 3 (mean = 2.02, median = 2)
proportion of cladograms that support basic monophyly = 0.88

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 39
(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 = 7.59, median = 9)
maximum number of subtaxa (genus) = 3 (mean = 2.9, median = 3)
proportion of cladograms that support rigorous monophyly = 0.72

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

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

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
101210324210
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
11119317315
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1313933319
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
111173188215
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Cumberlandia **            
     +--+                              
        |  +-- Pseudunio **               
        +--+                           
           +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
111173188217
  +-------- Gibbosula_rochechouartii
  |                                 
--+  +----- Gibbosula_laosensis     
  |  |                              
  +--+  +-- Cumberlandia **            
     |  |                           
     +--+-- Pseudunio **               
        |                           
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
111193186215
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Margaritifera **           
     +--+                              
        |  +-- Cumberlandia **            
        +--+                           
           +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
11119317317
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
101210358214
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
101210358212
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
101210358212
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10108324214
  +----------- Gibbosula_rochechouartii
--+                                    
  |  +-------- Gibbosula_laosensis     
  +--+                                 
     |  +----- Margaritifera **           
     +--+                              
        |  +-- Cumberlandia **            
        +--+                           
           +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1010833114
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
10108319218
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1010833118
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
1011931113
  +-------- Cumberlandia ** 
--+                      
  |  +----- Pseudunio **    
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
91083130114
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
91073130115
  +----- Cumberlandia ** 
  |                   
--+  +-- Gibbosulinae 
  |  |                
  +--+-- Pseudunio **    
     |                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
9108367116
  +-------- Cumberlandia ** 
--+                      
  |  +----- Gibbosulinae 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
9108321118
  +-------- Pseudunio **    
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
9108367112
  +-------- Cumberlandia ** 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
9108321118
  +-------- Pseudunio **    
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Gibbosulinae 
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
885310012
  +----------- Margaritifera_margaritifera **
  |                                       
  |     +----- Margaritifera_dahurica__1 **  
  |     |                                 
  +-----+----- Margaritifera_dahurica__2 **  
--+     |                                 
  |     |  +-- Pseudunio **                  
  |     +--+                              
  |        +-- Cumberlandia **               
  |                                       
  |  +-------- Margaritifera_falcata **      
  +--+                                    
     |  +----- Margaritifera_laevis **       
     +--+                                 
        |  +-- Margaritifera_middendorffi ** 
        +--+                              
           +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
88631004
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
88631004
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
88632009
  +----- Pseudunio_auricularius__1 **
  |                               
--+----- Pseudunio_auricularius__2 **
  |                               
  |  +-- Cumberlandia **             
  +--+                            
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
77535007
  +----- Margaritifera **
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
77531005
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
77434006
  +-- Pseudunio **    
  |                
--+-- Cumberlandia ** 
  |                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
773350010
  +-------- Margaritifera_dahurica **        
  |                                       
  +-------- Margaritifera_margaritifera__1 **
  |                                       
  +-------- Margaritifera_margaritifera__2 **
  |                                       
--+-------- Margaritifera_margaritifera__3 **
  |                                       
  |     +-- Pseudunio **                     
  +-----+                                 
  |     +-- Cumberlandia **                  
  |                                       
  |  +----- Margaritifera_falcata **         
  +--+                                    
     |  +-- Margaritifera_middendorffi **    
     +--+                                 
        +-- Margaritifera_marrianae **

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

iTtTbiriSoTroTroSiD
664385119
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Cumberlandia ** 
     +--+                
        +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
664335116
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
664335116
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
662320014
  +-------- 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
664330006
  +----- Cumberlandia ** 
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
664385119
  +-------- Gibbosulinae 
--+                      
  |  +----- Margaritifera **
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
553337117
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553368117
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553351117
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553390217
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553363217
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553368217
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
55335217
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553368217
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553383117
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
553390217
  +-------- Gibbosulinae 
--+                      
  |  +----- Cumberlandia ** 
  +--+                   
     |  +-- Pseudunio **    
     +--+                
        +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
4422340010
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
45011141110
  +-- Gibbosulinae               
  |                              
  +-- Margaritifera_dahurica **     
  |                              
--+-- Margaritifera_margaritifera **
  |                              
  +-- Margaritifera_laevis **       
  |                              
  +-- Margaritifera_middendorffi **

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

iTtTbiriSoTroTroSiD
442233008
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
4401320010
1

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

iTtTbiriSoTroTroSiD
331246119
  +----- Gibbosulinae 
--+                   
  |  +-- Cumberlandia ** 
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
31952160025
  +----------- Cumberlandia **       
  |                               
  +----------- Margaritifera **      
  |                               
  |  +-------- Gonideinae         
--+  |                            
  |  |     +-- Elliptio           
  |  |     |                      
  |  |     +-- Fusconaia          
  |  |     |                      
  +--+  +--+-- Amblemini          
     |  |  |                      
     |  |  +-- Pleurobema         
     |  |  |                      
     |  |  +-- Unionini           
     |  |                         
     |  |  +-- Pyganodon_cataracta
     +--+  |                      
        |  +-- Pyganodon_grandis  
        |  |                      
        |  +-- Utterbackia        
        |  |                      
        |  +-- Lampsilis_teres    
        |  |                      
        |  +-- Lampsilis_straminea
        +--+                      
           +-- Megalonaias        
           |                      
           +-- Obliquaria         
           |                      
           +-- Theliderma         
           |                      
           +-- Quadrula           
           |                      
           +-- Uniomerus

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

iTtTbiriSoTroTroSiD
331241009
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331235009
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331237009
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3312110010
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
32232190027
  +-------- 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        
        |                      
        +-- Aculamprotula      
        |                      
        +-- Unio

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

iTtTbiriSoTroTroSiD
33110009
1

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

iTtTbiriSoTroTroSiD
331243119
  +----- Gibbosulinae 
--+                   
  |  +-- Cumberlandia ** 
  +--+                
     +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
3313190011
  +----- Pseudunio **    
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
33131009
  +----- Cumberlandia ** 
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
331224009
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
32242190027
  +----------- 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

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

iTtTbiriSoTroTroSiD
331229009
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331330011
  +----- Margaritifera **
--+                   
  |  +-- Pseudunio **    
  +--+                
     +-- Cumberlandia **

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

iTtTbiriSoTroTroSiD
331245009
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331225009
  +-- Cumberlandia ** 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331241009
  +-- Pseudunio **    
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
331319009
  +----- Cumberlandia ** 
--+                   
  |  +-- Margaritifera **
  +--+                
     +-- Pseudunio **

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

iTtTbiriSoTroTroSiD
2201600--
1

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

iTtTbiriSoTroTroSiD
21501140025
  +-------------- Margaritifera_margaritifera__2 **
  |                                             
--+           +-- Margaritifera_margaritifera__1 **
  |  +--------+                                 
  |  |        +-- Margaritifera_falcata **         
  +--+                                          
     |  +-------- Unioninae                     
     |  |                                       
     +--+     +-- Pleurobemini                  
        |  +--+                                 
        |  |  +-- Gonideinae                    
        +--+                                    
           |  +-- Lampsilini                    
           +--+                                 
              +-- Quadrulini

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
22013400--
1

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

iTtTbiriSoTroTroSiD
22011700--
1

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

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

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

iTtTbiriSoTroTroSiD
2201800--
1

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

iTtTbiriSoTroTroSiD
22015100--
1

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
22011000--
1

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

iTtTbiriSoTroTroSiD
22017400--
1

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

iTtTbiriSoTroTroSiD
22015500--
1

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

iTtTbiriSoTroTroSiD
22012900--
1

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

iTtTbiriSoTroTroSiD
2201500--
1

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

iTtTbiriSoTroTroSiD
2201400--
1

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

iTtTbiriSoTroTroSiD
22015200--
1

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
22018900--
1

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

iTtTbiriSoTroTroSiD
2201400--
1

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

iTtTbiriSoTroTroSiD
22013500--
1

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

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

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

iTtTbiriSoTroTroSiD
22013600--
1

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
22011200--
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
22011700--
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
2201600--
1

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

iTtTbiriSoTroTroSiD
2201400--
1

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

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

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

iTtTbiriSoTroTroSiD
22015500--
1

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

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

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

iTtTbiriSoTroTroSiD
22011000--
1

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
220111900--
1

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

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

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

iTtTbiriSoTroTroSiD
22013100--
1

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

iTtTbiriSoTroTroSiD
2201700--
1

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
22012000--
1

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

iTtTbiriSoTroTroSiD
22011400--
1

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

iTtTbiriSoTroTroSiD
23402320042
  +-------- Cumberlandia **   
  |                        
  +-------- Margaritifera **  
--+                        
  |  +----- Unioninae      
  |  |                     
  |  |  +-- Ptychorhynchus 
  +--+  |                  
     |  +-- Chamberlainiini
     |  |                  
     +--+-- Lamprotulini   
        |                  
        +-- Sinosolenaia   
        |                  
        +-- Ambleminae

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

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

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

iTtTbiriSoTroTroSiD
22011200--
1

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

iTtTbiriSoTroTroSiD
23402320042
  +----------------- Cumberlandia **   
  |                                 
--+----------------- Margaritifera **  
  |                                 
  |  +-------------- Unioninae      
  +--+                              
     |  +----------- Sinosolenaia   
     +--+                           
        |  +-------- Lamprotulini   
        +--+                        
           |  +----- Chamberlainiini
           +--+                     
              |  +-- Ptychorhynchus 
              +--+                  
                 +-- Ambleminae

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
2201900--
1

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

iTtTbiriSoTroTroSiD
22013100--
1

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

iTtTbiriSoTroTroSiD
22011900--
1

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

iTtTbiriSoTroTroSiD
22012900--
1

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

iTtTbiriSoTroTroSiD
22018900--
1

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

iTtTbiriSoTroTroSiD
22011000--
1

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
22012600--
1

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

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

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

iTtTbiriSoTroTroSiD
22013500--
1

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

iTtTbiriSoTroTroSiD
2201600--
1

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

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

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

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

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

iTtTbiriSoTroTroSiD
22013600--
1

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

iTtTbiriSoTroTroSiD
22025111--
  +----- Gibbosulinae
--+                  
  |  +-- Pseudunio **   
  +--+               
     +-- Cumberlandia **

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

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

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

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

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

iTtTbiriSoTroTroSiD
11012200--
-- Pseudunio_marocanus **

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

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

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

iTtTbiriSoTroTroSiD
11011600--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11011400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011400--
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
1101500--
-- 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 **

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

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

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

iTtTbiriSoTroTroSiD
1101900--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1101900--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11015600--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11015500--
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
11013911--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11013911--
  +-- Gibbosulinae 
--+                
  +-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
11011100--
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

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

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

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011800--
-- Pseudunio_marocanus **

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

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

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
11011700--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
11011400--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101600--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101800--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011900--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101200--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11012800--
-- Margaritifera_falcata **

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

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

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

iTtTbiriSoTroTroSiD
11011000--
-- Pseudunio_auricularius **

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

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

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

iTtTbiriSoTroTroSiD
1101900--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
1101300--
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
11014811--
-- Margaritifera_dahurica **

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

iTtTbiriSoTroTroSiD
11011000--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

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

Graf, 2000 [Publ 2647], fig. 1

iTtTbiriSoTroTroSiD
11011600--
-- Margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011000--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11014211--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1101700--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011300--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011600--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011700--
-- Pseudunio_auricularius **

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

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

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

iTtTbiriSoTroTroSiD
1101900--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101700--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11012011--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
1101400--
-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
11011300--
-- Margaritifera_middendorffi **

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

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

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011400--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Margaritifera_hembeli **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
11011800--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
110110300--
-- Cumberlandia_monodonta **

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

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

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

iTtTbiriSoTroTroSiD
11013000--
-- Margaritifera_margaritifera **

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

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

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

iTtTbiriSoTroTroSiD
11011400--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
1101900--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11012011--
  +-- Margaritifera **
--+                
  +-- Gibbosulinae

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

iTtTbiriSoTroTroSiD
11011300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11012700--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011500--
-- Margaritifera_falcata **

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

iTtTbiriSoTroTroSiD
1101500--
-- Margaritifera_margaritifera **

Bashê, 2022 [Publ 4265], fig. 1

iTtTbiriSoTroTroSiD
1101500--
-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
1101300--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
11011000--
-- Margaritifera_margaritifera **

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

iTtTbiriSoTroTroSiD
1101700--
-- Cumberlandia_monodonta **

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

iTtTbiriSoTroTroSiD
11011300--
-- Cumberlandia_monodonta **
 
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