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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!

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Bivalvia | Unionoida | Unionoidea | Unionidae | Ambleminae

tribe QUADRULINI Ihering, 1901: cladomics
     5 genera, 1 subtribe

Cladomics Summary

number of species = 26 (46 total)
number of subtaxa (rank: subtribe) = 6
analyzed in 194 cladograms in 118 publications
maximum number of terminals in an analysis = 22 (mean = 2.59, median = 1)
maximum number of subtaxa (subtribe) = 3 (mean = 1.36, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 87
maximum number of included terminals = 22, (mean = 4.55 median = 2)
maximum number of subtaxa (subtribe) = 3 (mean = 1.8, median = 2)
proportion of cladograms that support basic monophyly = 0.83

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 50
(Rigorous monophyly refers to cladograms with terminal species representing more than one subtaxon (subtribe) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 22 (mean = 5.74, median = 4)
maximum number of subtaxa (subtribe) = 3 (mean = 2.34, median = 2)
proportion of cladograms that support rigorous monophyly = 0.76


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


Neemuchwala et al., 2023 [Publ 4292], fig. 2

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
22222035440
        +----- Megalonaias ** 
  +-----+                  
  |     |  +-- Uniomerus **   
  |     +--+               
--+        +-- Quadrulina **  
  |                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Lampsilini  
        +--+               
           +-- Popenaiadini

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
1111839179420
           +----- Quadrulina **  
  +--------+                  
  |        |  +-- Megalonaias ** 
  |        +--+               
--+           +-- Uniomerus **   
  |                           
  |  +----------- Pleurobemini
  |  |                        
  +--+        +-- Plectomerus 
     |  +-----+               
     |  |     +-- Reginaia    
     +--+                     
        |  +----- Lampsilini  
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Amblema     

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

iTtTbiriSoTroTroSiD
1925233106327
  +----------- Megalonaias ** 
  |                        
--+        +-- Uniomerus **   
  |  +-----+               
  |  |     +-- Quadrulina **  
  +--+                     
     |  +----- Pleurobemini
     +--+                  
        |  +-- Lampsilini  
        +--+               
           +-- Amblemini   

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

iTtTbiriSoTroTroSiD
1818153106323
  +-------------- Amblemini   
--+                           
  |  +----------- Lampsilini  
  +--+                        
     |  +-------- Pleurobemini
     +--+                     
        |  +----- Quadrulina **  
        +--+                  
           |  +-- Megalonaias ** 
           +--+               
              +-- Uniomerus **   

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

iTtTbiriSoTroTroSiD
1616143106322
     +----- Amblemini   
  +--+                  
  |  |  +-- Pleurobemini
  |  +--+               
--+     +-- Lampsilini  
  |                     
  |  +----- Megalonaias ** 
  +--+                  
     |  +-- Uniomerus **   
     +--+               
        +-- Quadrulina **  

Johnson et al., 2018 [Publ 3844], fig. 2

iTtTbiriSoTroTroSiD
141412322220
  +----------- Amblemini   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Megalonaias ** 
     +--+                  
        |  +-- Uniomerus **   
        +--+               
           +-- Quadrulina **  

Szumowski et al., 2012 [Publ 3569], fig. 5

iTtTbiriSoTroTroSiD
141412200020
  +----- Megalonaias **
  |                 
--+----- Theliderma ** 
  |                 
  |  +-- Quadrula **   
  +--+              
     +-- Pustulosa **  

Serb et al., 2003 [Publ 2659], fig. 3

iTtTbiriSoTroTroSiD
14141221515322
  +-------------------------- Lampsilis   
  |                                       
--+  +----------------------- Potamilus   
  |  |                                    
  |  |                 +----- Medionidus  
  +--+  +--------------+                  
     |  |              |  +-- Dromus      
     |  |              +--+               
     +--+                 +-- Cyprogenia  
        |                                 
        |  +----------------- Obliquaria  
        |  |                              
        +--+  +-------------- Amblema     
           |  |                           
           |  |        +----- Reginaia    
           +--+  +-----+                  
              |  |     |  +-- Elliptoideus
              |  |     +--+               
              |  |        +-- Plectomerus 
              +--+                        
                 |        +-- Megalonaias ** 
                 |  +-----+               
                 |  |     +-- Quadrulina **  
                 +--+                     
                    |  +----- Hemistena   
                    +--+                  
                       |  +-- Elliptio    
                       +--+               
                          +-- Fusconaia   

Serb et al., 2003 [Publ 2659], fig. 2

iTtTbiriSoTroTroSiD
14141121515317
  +----------- Lampsilis   
  |                        
  |  +-------- Obliquaria  
  |  |                     
  |  +-------- Amblema     
  |  |                     
--+  +-------- Reginaia    
  |  |                     
  |  +-------- Potamilus   
  |  |                     
  |  |     +-- Elliptoideus
  |  +-----+               
  |  |     +-- Plectomerus 
  +--+                     
     |     +-- Megalonaias ** 
     +-----+               
     |     +-- Quadrulina **  
     |                     
     |  +----- Medionidus  
     +--+                  
     |  |  +-- Dromus      
     |  +--+               
     |     +-- Cyprogenia  
     |                     
     |  +----- Hemistena   
     +--+                  
        |  +-- Elliptio    
        +--+               
           +-- Fusconaia   

Campbell & Lydeard, 2012 [Publ 3309], fig. 3

iTtTbiriSoTroTroSiD
1111826262422
                                   +-- Uniomerus **             
  +--------------------------------+                         
  |                                +-- Quadrulina **            
  |                                                          
--+  +-------------------------------- Pleurobemini          
  |  |                                                       
  |  |                             +-- Amblema               
  +--+  +--------------------------+                         
     |  |                          +-- Reginaia              
     |  |                                                    
     +--+  +-------------------------- Popenaiadini          
        |  |                                                 
        |  |                       +-- Glebula               
        +--+  +--------------------+                         
           |  |                    +-- Paetulunio            
           |  |                                              
           +--+                    +-- Toxolasma             
              |  +-----------------+                         
              |  |                 +-- Plectomerus           
              |  |                                           
              +--+  +----------------- Obliquaria            
                 |  |                                        
                 |  |           +----- Potamilus             
                 +--+  +--------+                            
                    |  |        |  +-- Dromus                
                    |  |        +--+                         
                    +--+           +-- Lemiox                
                       |                                     
                       |  +----------- Lampsilis_ovata       
                       |  |                                  
                       +--+  +-------- Lampsilis_virescens   
                          |  |                               
                          +--+     +-- Epioblasma            
                             |  +--+                         
                             |  |  +-- Obovaria_arkansasensis
                             +--+                            
                                |  +-- Obovaria_olivaria     
                                +--+                         
                                   +-- Venustaconcha         

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

iTtTbiriSoTroTroSiD
1111839179420
                       +----- Megalonaias **                   
  +--------------------+                                    
  |                    |  +-- Uniomerus **                     
  |                    +--+                                 
  |                       +-- Quadrulina **                    
  |                                                         
--+  +----------------------- Pleurobemini                  
  |  |                                                      
  |  |                 +----- Popenaiadini                  
  |  |  +--------------+                                    
  |  |  |              |  +-- Amblema                       
  +--+  |              +--+                                 
     |  |                 +-- Reginaia                      
     |  |                                                   
     |  |                 +-- Plectomerus                   
     |  |  +--------------+                                 
     +--+  |              +-- Toxolasma                     
        |  |                                                
        |  |           +----- Obliquaria                    
        |  |  +--------+                                    
        |  |  |        |  +-- Cyrtonaias                    
        |  |  |        +--+                                 
        |  |  |           +-- Glebula                       
        |  |  |                                             
        +--+  |  +----------- Sagittunio                    
           |  |  |                                          
           |  |  +----------- Leaunio                       
           |  |  |                                          
           |  |  +----------- Potamilus                     
           |  |  |                                          
           |  |  +----------- Hamiota                       
           |  |  |                                          
           |  |  |        +-- Ellipsaria                    
           +--+  +--------+                                 
              |  |        +-- Truncilla                     
              |  |                                          
              |  |     +----- Lampsilis_teres               
              |  +-----+                                    
              |  |     |  +-- Ortmanniana_ligamentina       
              |  |     +--+                                 
              |  |        +-- Lampsilis_siliquoidea         
              |  |                                          
              |  |     +----- Villosa                       
              |  +-----+                                    
              |  |     |  +-- Ligumia                       
              |  |     +--+                                 
              |  |        +-- Cambarunio                    
              |  |                                          
              +--+  +-------- Ortmanniana_pectorosa         
                 +--+                                       
                 |  |  +----- Lampsilis_ornata              
                 |  +--+                                    
                 |     |  +-- Lampsilis_cardium             
                 |     +--+                                 
                 |        +-- Lampsilis_ovata               
                 |                                          
                 |  +-------- Obovaria_olivaria             
                 |  |                                       
                 |  +-------- Venustaconcha_ellipsiformis__2
                 +--+                                       
                 |  +-------- Epioblasma                    
                 |  |                                       
                 |  |  +----- Obovaria_unicolor             
                 |  +--+                                    
                 |     |  +-- Obovaria_arkansasensis        
                 |     +--+                                 
                 |        +-- Obovaria_subrotunda           
                 |                                          
                 |        +-- Ptychobranchus                
                 |  +-----+                                 
                 |  |     +-- Venustaconcha_ellipsiformis__1
                 |  |                                       
                 +--+  +----- Medionidus_conradicus         
                    |  |                                    
                    |  |  +-- Lemiox                        
                    +--+--+                                 
                       |  +-- Medionidus_acutissimus        
                       |                                    
                       |  +-- Dromus                        
                       +--+                                 
                          +-- Cyprogenia                    

Campbell & Lydeard, 2012 [Publ 3309], fig. 1-2

iTtTbiriSoTroTroSiD
1111926262421
              +-- Uniomerus **   
  +-----------+               
  |           +-- Quadrulina **  
--+                           
  |  +----------- Pleurobemini
  |  |                        
  +--+        +-- Plectomerus 
     |  +-----+               
     |  |     +-- Reginaia    
     +--+                     
        |  +----- Lampsilini  
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Amblema     

Boyer et al., 2011 [Publ 2718], fig. 4

iTtTbiriSoTroTroSiD
7292523122341
  +-------------------------------- Quadrulina **           
  |                                                      
--+  +----------------------------- Megalonaias **          
  |  |                                                   
  +--+  +-------------------------- Toxolasma            
     |  |                                                
     +--+  +----------------------- Amblema              
        |  |                                             
        |  |  +-------------------- Pleurobemini         
        +--+  |                                          
           |  |                 +-- Reginaia             
           |  +-----------------+                        
           +--+                 +-- Obliquaria           
              |                                          
              |  +----------------- Potamilus            
              |  |                                       
              |  |              +-- Ellipsaria           
              +--+  +-----------+                        
                 |  |           +-- Truncilla            
                 |  |                                    
                 +--+  +----------- Lampsilis_cardium    
                    |  |                                 
                    |  |     +----- Ligumia              
                    |  +-----+                           
                    +--+     |  +-- Obovaria             
                       |     +--+                        
                       |        +-- Epioblasma           
                       |                                 
                       |  +-------- Lampsilis_teres      
                       +--+                              
                          |  +----- Lampsilis_siliquoidea
                          +--+                           
                             |  +-- Ortmanniana          
                             +--+                        
                                +-- Lampsilis_higginsii  

Boyer et al., 2011 [Publ 2718], fig. 3

iTtTbiriSoTroTroSiD
77522822323
  +----------- Reginaia    
  |                        
--+        +-- Amblema     
  |  +-----+               
  |  |     +-- Lampsilini  
  +--+                     
     |  +----- Pleurobemini
     +--+                  
        |  +-- Megalonaias ** 
        +--+               
           +-- Quadrulina **  

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

iTtTbiriSoTroTroSiD
77229775419
  +-------- Reginaia           
  |                            
  +-------- Amblema            
  |                            
  |  +----- Pleurobemini       
  +--+                         
  |  |  +-- Uniomerus **          
  |  +--+                      
--+     +-- Quadrulina **         
  |                            
  |  +----- Popenaiadini       
  |  |                         
  |  |  +-- Dromus             
  |  |  |                      
  |  |  +-- Epioblasma         
  |  |  |                      
  +--+  +-- Glebula            
     |  |                      
     |  +-- Lampsilis_ovata    
     |  |                      
     |  +-- Lampsilis_virescens
     |  |                      
     |  +-- Lemiox             
     +--+                      
        +-- Obliquaria         
        |                      
        +-- Obovaria_olivaria  
        |                      
        +-- Obovaria_unicolor  
        |                      
        +-- Plectomerus        
        |                      
        +-- Potamilus_alatus   
        |                      
        +-- Potamilus_capax    

Zhou et al., 2007 [Publ 2658], fig. 2

iTtTbiriSoTroTroSiD
7751110023
1

Campbell & Lydeard, 2012 [Publ 3308], fig. 3

iTtTbiriSoTroTroSiD
77523636421
                                +-- Uniomerus **           
  +-----------------------------+                       
  |                             +-- Quadrulina **          
  |                                                     
  |                 +-------------- Elliptoideus        
  |  +--------------+                                   
  |  |              |  +----------- Plectomerus         
--+  |              +--+                                
  |  |                 |  +-------- Obovaria_unicolor   
  |  |                 +--+                             
  |  |                    |  +----- Obovaria_olivaria   
  |  |                    +--+                          
  |  |                       |  +-- Lampsilis           
  +--+                       +--+                       
     |                          +-- Lemiox              
     |                                                  
     |                    +-------- Cyrtonaias          
     |  +-----------------+                             
     |  |                 |  +----- Reginaia            
     |  |                 +--+                          
     |  |                    |  +-- Popenaiadini        
     +--+                    +--+                       
        |                       +-- Amblema             
        |                                               
        |  +----------------------- Elliptio            
        |  |                                            
        +--+  +-------------------- Eurynia             
           |  |                                         
           |  |  +----------------- Parvaspina          
           +--+  |                                      
              |  |              +-- Hemistena           
              |  +--------------+                       
              +--+              +-- Plethobasus         
                 |                                      
                 |  +-------------- Pleuronaia          
                 |  |                                   
                 |  |           +-- Pleurobema_cordatum 
                 +--+  +--------+                       
                    |  |        +-- Fusconaia           
                    |  |                                
                    +--+        +-- Pleurobema_stabile  
                       |  +-----+                       
                       |  |     +-- Pleurobema_sintoxia 
                       +--+                             
                          |  +----- Pleurobema_decisum  
                          +--+                          
                             |  +-- Pleurobema_clava    
                             +--+                       
                                +-- Pleurobema_pyriforme

Campbell & Lydeard, 2012 [Publ 3308], fig. 2

iTtTbiriSoTroTroSiD
66413636422
  +-------------------- Quadrulina **  
  |                                 
--+  +----------------- Pleurobemini
  |  |                              
  +--+              +-- Reginaia    
     |  +-----------+               
     |  |           +-- Amblema     
     +--+                           
        |  +----------- Popenaiadini
        |  |                        
        +--+  +-------- Cyrtonaias  
           |  |                     
           +--+-------- Plectomerus 
              |                     
              |  +----- Lemiox      
              +--+                  
                 |  +-- Lampsilis   
                 +--+               
                    +-- Obovaria    

Pfeiffer et al., 2016 [Publ 3660], fig. 2

iTtTbiriSoTroTroSiD
66423231322
  +----------------------------- Amblemini            
  |                                                   
--+  +-------------------------- Lampsilini           
  |  |                                                
  +--+  +----------------------- Elliptoideus         
     |  |                                             
     |  |                    +-- Uniomerus **            
     +--+  +-----------------+                        
        |  |                 +-- Quadrulina **           
        |  |                                          
        +--+  +----------------- Elliptio             
           |  |                                       
           |  |  +-------------- Parvaspina           
           +--+  |                                    
              |  +-------------- Plethobasus          
              |  |                                    
              |  |           +-- Hemistena            
              +--+-----------+                        
                 |           +-- Eurynia              
                 |                                    
                 |  +----------- Fusconaia            
                 |  |                                 
                 |  |     +----- Pleurobema_riddellii 
                 +--+  +--+                           
                    |  |  |  +-- Pleurobema_sintoxia  
                    |  |  +--+                        
                    +--+     +-- Pleurobema_cordatum  
                       |                              
                       |  +----- Pleuronaia           
                       |  |                           
                       +--+  +-- Pleurobema_pyriforme 
                          |  |                        
                          +--+-- Pleurobema_strodeanum
                             |                        
                             +-- Pleurobema_clava     

Campbell & Lydeard, 2012 [Publ 3308], fig. 1

iTtTbiriSoTroTroSiD
66413636422
  +-------------------- Quadrulina **  
  |                                 
--+  +----------------- Pleurobemini
  |  |                              
  |  |              +-- Popenaiadini
  +--+  +-----------+               
     |  |           +-- Amblema     
     |  |                           
     +--+           +-- Cyrtonaias  
        |  +--------+               
        |  |        +-- Reginaia    
        +--+                        
           |  +-------- Plectomerus 
           +--+                     
              |  +----- Lemiox      
              +--+                  
                 |  +-- Lampsilis   
                 +--+               
                    +-- Obovaria    

Campbell & Lydeard, 2012 [Publ 3308], fig. 4

iTtTbiriSoTroTroSiD
66413636422
  +----------------- Quadrulina **  
  |                              
--+  +-------------- Pleurobemini
  |  |                           
  |  |           +-- Popenaiadini
  +--+  +--------+               
     |  |        +-- Amblema     
     |  |                        
     +--+  +-------- Plectomerus 
        |  |                     
        |  |     +-- Cyrtonaias  
        +--+-----+               
           |     +-- Reginaia    
           |                     
           |  +----- Lemiox      
           +--+                  
              |  +-- Lampsilis   
              +--+               
                 +-- Obovaria    

Pfeiffer et al., 2019 [Publ 3866], fig. 1

iTtTbiriSoTroTroSiD
55332625419
        +----- Megalonaias ** 
  +-----+                  
  |     |  +-- Uniomerus **   
  |     +--+               
--+        +-- Quadrulina **  
  |                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
41703187333
  +-- Gonideini          
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias **        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus **          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Aculamprotula      
  |                      
  +-- Unio               

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

iTtTbiriSoTroTroSiD
4181632414334
              +-- Megalonaias ** 
  +-----------+               
  |           +-- Quadrulina **  
  |                           
--+        +----- Amblema     
  |  +-----+                  
  |  |     |  +-- Plectomerus 
  |  |     +--+               
  +--+        +-- Lampsilini  
     |                        
     |  +-------- Elliptoideus
     +--+                     
        |  +----- Uniomerus **   
        +--+                  
           |  +-- Eurynia     
           +--+               
              +-- Pleurobema  

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

iTtTbiriSoTroTroSiD
4181432314334
                          +-- Megalonaias **         
  +-----------------------+                       
  |                       +-- Quadrulina **          
  |                                               
  |                    +----- Elliptoideus        
--+                    |                          
  |  +-----------------+  +-- Eurynia             
  |  |                 |  |                       
  |  |                 +--+-- Pleurobema          
  |  |                    |                       
  +--+                    +-- Uniomerus **           
     |                                            
     |  +-------------------- Amblema             
     |  |                                         
     +--+                 +-- Plectomerus         
        |  +--------------+                       
        |  |              +-- Obliquaria          
        +--+                                      
           |  +-------------- Potamilus_fragilis  
           |  |                                   
           +--+  +----------- Ptychobranchus      
              |  |                                
              +--+  +-------- Medionidus          
                 |  |                             
                 +--+  +----- Potamilus_purpuratus
                    |  |                          
                    +--+----- Lampsilis           
                       |                          
                       |  +-- Ellipsaria          
                       +--+                       
                          +-- Villosa             

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

iTtTbiriSoTroTroSiD
442319721320
                       +----- Quadrulina **        
  +--------------------+                        
  |                    |  +-- Megalonaias **       
  |                    +--+                     
--+                       +-- Uniomerus **         
  |                                             
  |  +----------------------- Pleurobemini      
  |  |                                          
  +--+  +-------------------- Amblema           
     |  |                                       
     |  |              +----- Toxolasma         
     +--+  +-----------+                        
        |  |           |  +-- Obliquaria        
        |  |           +--+                     
        |  |              +-- Plectomerus       
        +--+                                    
           |  +-------------- Potamilus_leptodon
           |  |                                 
           |  |  +----------- Potamilus_alatus  
           +--+  |                              
              |  |  +-------- Venustaconcha     
              |  |  |                           
              +--+  +-------- Leaunio           
                 |  |                           
                 |  |     +-- Ellipsaria        
                 |  +-----+                     
                 +--+     +-- Truncilla         
                    |                           
                    |     +-- Medionidus        
                    +-----+                     
                    |     +-- Ptychobranchus    
                    |                           
                    |  +----- Cambarunio        
                    +--+                        
                       |  +-- Ortmanniana       
                       +--+                     
                          +-- Lampsilis         

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

iTtTbiriSoTroTroSiD
4128733197213145
  +----------- Chamberlainia        
  |                                 
  +----------- Megalonaias **          
  |                                 
  +----------- Uniomerus **            
  |                                 
  +----------- Quadrulina **           
  |                                 
  +----------- Sinohyriopsis        
  |                                 
  +----------- Lamprotulini         
  |                                 
  +----------- Gonideini            
  |                                 
  +----------- Pseudodontini        
  |                                 
  |        +-- Rectidentini         
  +--------+                        
  |        +-- Contradentini        
  |                                 
  |  +-------- Amblema              
  |  |                              
  |  +-------- Toxolasma            
  |  |                              
  |  +-------- Pleurobemini         
  |  |                              
  |  |     +-- Obliquaria           
  |  +-----+                        
  |  |     +-- Plectomerus          
  +--+                              
  |  |  +----- Ellipsaria           
  |  |  |                           
  |  |  +----- Potamilus_leptodon   
  |  |  |                           
  |  |  +----- Medionidus           
  |  |  |                           
  |  |  +----- Potamilus_alatus     
--+  |  |                           
  |  |  +----- Ptychobranchus       
  |  +--+                           
  |     +----- Truncilla            
  |     |                           
  |     +----- Venustaconcha        
  |     |                           
  |     +----- Cambarunio           
  |     |                           
  |     +----- Leaunio              
  |     |                           
  |     |  +-- Ortmanniana          
  |     +--+                        
  |        +-- Lampsilis            
  |                                 
  |  +-------- Acuticostini         
  |  |                              
  |  +-------- Alasmidonta          
  |  |                              
  |  +-------- Anodonta_anatina     
  |  |                              
  |  +-------- Anemina              
  |  |                              
  |  +-------- Anodonta_cygnea      
  |  |                              
  |  +-------- Cristaria            
  |  |                              
  |  +-------- Anodonta_nuttalliana 
  |  |                              
  |  +-------- Lasmigona            
  |  |                              
  |  +-------- Lepidodesmini        
  |  |                              
  |  +-------- Pseudanodonta        
  |  |                              
  |  +-------- Pyganodon            
  |  |                              
  +--+-------- Simpsonaias          
     |                              
     +-------- Strophitus           
     |                              
     +-------- Utterbackia          
     |                              
     +-------- Lanceolariini        
     |                              
     +-------- Schistodesmus        
     |                              
     +-------- Unio                 
     |                              
     +-------- Aculamprotula        
     |                              
     |     +-- Pletholophus         
     +-----+                        
     |     +-- Sinanodonta          
     |                              
     |  +----- Cuneopsis_celtiformis
     |  |                           
     |  +----- Tchangsinaia         
     |  |                           
     +--+----- Cuneopsis_rufescens  
        |                           
        +----- Nodularia            
        |                           
        |  +-- Cuneopsis_szechenyii 
        +--+                        
           +-- Cuneopsis_heudei     

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

iTtTbiriSoTroTroSiD
4212032917338
                          +-- Megalonaias **          
  +-----------------------+                        
  |                       +-- Quadrulina **           
  |                                                
  |              +----------- Elliptoideus         
  |  +-----------+                                 
--+  |           |  +-------- Pleurobema           
  |  |           +--+                              
  |  |              |  +----- Uniomerus **            
  |  |              +--+                           
  |  |                 |  +-- Fusconaia            
  +--+                 +--+                        
     |                    +-- Eurynia              
     |                                             
     |  +-------------------- Amblema              
     |  |                                          
     |  |              +----- Toxolasma            
     |  |  +-----------+                           
     +--+  |           |  +-- Plectomerus          
        |  |           +--+                        
        |  |              +-- Obliquaria           
        |  |                                       
        +--+           +----- Ptychobranchus       
           |  +--------+                           
           |  |        |  +-- Potamilus_fragilis   
           |  |        +--+                        
           |  |           +-- Medionidus           
           +--+                                    
              |           +-- Villosa              
              |  +--------+                        
              |  |        +-- Ellipsaria           
              +--+                                 
                 |  +-------- Lampsilis_teres      
                 +--+                              
                    |  +----- Lampsilis_siliquoidea
                    +--+                           
                       |  +-- Potamilus_purpuratus 
                       +--+                        
                          +-- Potamilus_alatus     

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

iTtTbiriSoTroTroSiD
4323123314349
  +-------------- Quadrulina **     
  |                              
  |        +----- Lamprotulini   
--+  +-----+                     
  |  |     |  +-- Chamberlainiini
  |  |     +--+                  
  +--+        +-- Anodontini     
     |                           
     |  +-------- Megalonaias **    
     +--+                        
        |  +----- Pleurobemini   
        +--+                     
           |  +-- Amblemini      
           +--+                  
              +-- Lampsilini     

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

iTtTbiriSoTroTroSiD
442319521320
                                +----- Quadrulina **        
  +-----------------------------+                        
  |                             |  +-- Megalonaias **       
  |                             +--+                     
--+                                +-- Uniomerus **         
  |                                                      
  |  +-------------------------------- Pleurobemini      
  |  |                                                   
  +--+  +----------------------------- Amblema           
     |  |                                                
     +--+  +-------------------------- Toxolasma         
        |  |                                             
        +--+                       +-- Obliquaria        
           |  +--------------------+                     
           |  |                    +-- Plectomerus       
           +--+                                          
              |  +-------------------- Potamilus_leptodon
              |  |                                       
              +--+  +----------------- Potamilus_alatus  
                 |  |                                    
                 |  |              +-- Medionidus        
                 +--+  +-----------+                     
                    |  |           +-- Ptychobranchus    
                    |  |                                 
                    +--+           +-- Ellipsaria        
                       |  +--------+                     
                       |  |        +-- Truncilla         
                       +--+                              
                          |  +-------- Leaunio           
                          |  |                           
                          +--+     +-- Venustaconcha     
                             |  +--+                     
                             |  |  +-- Cambarunio        
                             +--+                        
                                |  +-- Ortmanniana       
                                +--+                     
                                   +-- Lampsilis         

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

iTtTbiriSoTroTroSiD
41703188333
  +-- Gonideini          
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias **        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus **          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Pleurobema         
  |                      
  +-- Unionini           

Pfeiffer et al., 2018 [Publ 3835], fig. 2

iTtTbiriSoTroTroSiD
442300022
  +----- Quadrulina ** 
--+                 
  |  +-- Uniomerus **  
  +--+              
     +-- Megalonaias **

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

iTtTbiriSoTroTroSiD
41003157326
  +-- Pyganodon_cataracta
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Lampsilis_straminea
--+                      
  +-- Megalonaias **        
  |                      
  +-- Obliquaria         
  |                      
  +-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus **          

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

iTtTbiriSoTroTroSiD
3301119320
  +-------- Amblemini   
--+                     
  |  +----- Lampsilini  
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
3311119319
1

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

iTtTbiriSoTroTroSiD
3311119319
  +-- Quadrulina **  
--+               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
33123514319
  +----- Pleurobemini
--+                  
  |  +-- Megalonaias ** 
  +--+               
     +-- Quadrulina **  

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

iTtTbiriSoTroTroSiD
3301119320
  +-------- Lampsilini  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
220290154--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
22025283--
        +-- Quadrulina **  
  +-----+               
  |     +-- Uniomerus **   
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Popenaiadini
     +--+               
        +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
22011072--
  +-------- Quadrulina ** 
--+                    
  |  +----- Plectomerus
  +--+                 
     |  +-- Amblema    
     +--+              
        +-- Lampsilini 

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

iTtTbiriSoTroTroSiD
220217124--
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
2201211--
1

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

iTtTbiriSoTroTroSiD
220272134--
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220251104--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220144112--
                 +----- Pleurobemini  
  +--------------+                    
  |              |  +-- Obliquaria    
  |              +--+                 
--+                 +-- Quadrulina **    
  |                                   
  |  +----------------- Ptychobranchus
  +--+                                
     |  +-------------- Truncilla     
     +--+                             
        |  +----------- Ortmanniana   
        +--+                          
           |  +-------- Lampsilis     
           +--+                       
              |  +----- Cambarunio    
              +--+                    
                 |  +-- Ligumia       
                 +--+                 
                    +-- Epioblasma    

Chapman et al., 2008 [Publ 2667], fig. 1

iTtTbiriSoTroTroSiD
220119184--
  +-------------- Quadrulina **  
--+                           
  |  +----------- Pleurobemini
  +--+                        
     |  +-------- Amblema     
     +--+                     
        |  +----- Plectomerus 
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
22011442--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
2201972--
  +----------------- Glebula            
  |                                     
--+              +-- Amblema            
  |  +-----------+                      
  |  |           +-- Quadrulina **         
  +--+                                  
     |  +----------- Plectomerus        
     +--+                               
        |  +-------- Leaunio            
        +--+                            
           |  +----- Lampsilis_straminea
           +--+                         
              |  +-- Lampsilis_teres    
              +--+                      
                 +-- Potamilus          

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

iTtTbiriSoTroTroSiD
220144113--
           +-- Obliquaria    
  +--------+                 
  |        +-- Quadrulina **    
  |                          
--+  +-------- Amblemini     
  |  |                       
  |  +-------- Pleurobemini  
  +--+                       
     |  +----- Truncilla     
     |  |                    
     |  +----- Ptychobranchus
     +--+                    
        |  +-- Ortmanniana   
        |  |                 
        |  +-- Cambarunio    
        +--+                 
           +-- Ligumia       
           |                 
           +-- Epioblasma    

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

iTtTbiriSoTroTroSiD
22025482--
     +-- Quadrulina **  
  +--+               
  |  +-- Uniomerus **   
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220294154--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

Freeman et al., 2025 [Publ 4443], fig. 4

iTtTbiriSoTroTroSiD
220110104--
     +----- Amblemini   
  +--+                  
  |  |  +-- Pleurobemini
  |  +--+               
  |     +-- Quadrulina **  
--+                     
  |  +----- Obliquaria  
  |  |                  
  |  +----- Toxolasma   
  +--+                  
     +----- Popenaiadini
     |                  
     |  +-- Glebula     
     +--+               
        +-- Cyrtonaias  

Porto-Hannes et al., 2021 [Publ 4227], fig. 4

iTtTbiriSoTroTroSiD
2110199329
  +-------- Quadrula **    
  |                     
--+-------- Pustulosa **   
  |                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220145123--
              +-------- Amblemini     
  +-----------+                       
  |           |  +----- Pleurobemini  
  |           +--+                    
  |              |  +-- Obliquaria    
--+              +--+                 
  |                 +-- Quadrulina **    
  |                                   
  |  +----------------- Ptychobranchus
  +--+                                
     |  +-------------- Truncilla     
     +--+                             
        |  +----------- Ortmanniana   
        +--+                          
           |  +-------- Lampsilis     
           +--+                       
              |  +----- Cambarunio    
              +--+                    
                 |  +-- Ligumia       
                 +--+                 
                    +-- Epioblasma    

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

iTtTbiriSoTroTroSiD
2201211--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
220255134--
        +-- Quadrulina **  
  +-----+               
  |     +-- Uniomerus **   
  |                     
--+     +-- Pleurobemini
  |  +--+               
  |  |  +-- Amblemini   
  +--+                  
     |  +-- Popenaiadini
     +--+               
        +-- Lampsilini  

Freeman et al., 2025 [Publ 4443], fig. 5

iTtTbiriSoTroTroSiD
2201994--
        +-- Popenaiadini
  +-----+               
  |     +-- Lampsilini  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Pleurobemini
     +--+               
        +-- Quadrulina **  

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

iTtTbiriSoTroTroSiD
214023212332
  +----------------------------- Megalonaias **         
  |                                                  
  |                       +----- Elliptoideus        
--+  +--------------------+                          
  |  |                    |  +-- Uniomerus **           
  |  |                    +--+                       
  +--+                       +-- Pleurobema          
     |                                               
     |  +----------------------- Amblema             
     |  |                                            
     +--+                    +-- Obliquaria          
        |  +-----------------+                       
        |  |                 +-- Plectomerus         
        +--+                                         
           |  +----------------- Medionidus          
           +--+                                      
              |  +-------------- Potamilus_fragilis  
              +--+                                   
                 |  +----------- Ptychobranchus      
                 +--+                                
                    |  +-------- Potamilus_purpuratus
                    +--+                             
                       |  +----- Lampsilis           
                       +--+                          
                          |  +-- Villosa             
                          +--+                       
                             +-- Ellipsaria          

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

iTtTbiriSoTroTroSiD
22017473--
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
220294154--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

Chapman et al., 2008 [Publ 2744], fig. 2

iTtTbiriSoTroTroSiD
220119184--
  +-------------- Quadrulina **  
--+                           
  |  +----------- Pleurobemini
  +--+                        
     |  +-------- Amblema     
     +--+                     
        |  +----- Plectomerus 
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220113773--
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

Freeman et al., 2025 [Publ 4443], fig. 3

iTtTbiriSoTroTroSiD
220110104--
        +-- Popenaiadini
  +-----+               
  |     +-- Lampsilini  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Pleurobemini
     +--+               
        +-- Quadrulina **  

Inoue et al., 2020 [Publ 3941], fig. 3

iTtTbiriSoTroTroSiD
220140393--
  +-------- Quadrulina **  
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220255134--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220272134--
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
22025292--
     +-- Quadrulina **  
  +--+               
  |  +-- Uniomerus **   
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Lampsilini  

Burlakova et al., 2019 [Publ 3899], fig. 1

iTtTbiriSoTroTroSiD
220231314--
                          +-- Quadrulina **           
  +-----------------------+                        
  |                       +-- Uniomerus **            
  |                                                
  |  +----------------------- Plectomerus          
--+  |                                             
  |  |  +-------------------- Popenaiadini         
  |  |  |                                          
  |  |  +-------------------- Pleurobemini         
  +--+  |                                          
     |  |                 +-- Cyrtonaias           
     |  |              +--+                        
     |  |              |  +-- Glebula              
     |  +--------------+                           
     |  |              |  +-- Reginaia             
     +--+              +--+                        
        |                 +-- Obliquaria           
        |                                          
        |  +----------------- Toxolasma            
        |  |                                       
        |  |  +-------------- Potamilus            
        |  |  |                                    
        |  |  |           +-- Ellipsaria           
        |  |  +-----------+                        
        +--+  |           +-- Truncilla            
           |  |                                    
           |  |        +----- Medionidus           
           |  +--------+                           
           |  |        |  +-- Lemiox               
           |  |        +--+                        
           +--+           +-- Dromus               
              |                                    
              |  +----------- Cambarunio           
              |  |                                 
              |  +----------- Leaunio              
              |  |                                 
              |  +----------- Villosa              
              |  |                                 
              |  |        +-- Sagittunio           
              |  +--------+                        
              +--+        +-- Lampsilis_ovata      
                 |                                 
                 |     +----- Venustaconcha        
                 +-----+                           
                 |     |  +-- Epioblasma           
                 |     +--+                        
                 |        +-- Obovaria             
                 |                                 
                 |  +-------- Lampsilis_teres      
                 +--+                              
                    |  +----- Lampsilis_siliquoidea
                    +--+                           
                       |  +-- Lampsilis_hydiana    
                       +--+                        
                          +-- Lampsilis_straminea  

Pieri et al., 2018 [Publ 3867], fig. 3

iTtTbiriSoTroTroSiD
220118171--
  +-- Quadrulina **  
--+               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
2201211--
1

Mejia-Trujillo & Smith, 2022 [Publ 4274], fig. 2-F

iTtTbiriSoTroTroSiD
220213134--
     +-------------- Toxolasma            
  +--+                                    
  |  |  +----------- Popenaiadini         
  |  +--+                                 
  |     |  +-------- Amblemini            
  |     +--+                              
  |        |  +----- Pleurobemini         
--+        +--+                           
  |           |  +-- Quadrulina **           
  |           +--+                        
  |              +-- Uniomerus **            
  |                                       
  |     +----------- Potamilus            
  |     |                                 
  +-----+        +-- Lampsilis_cardium    
        |  +-----+                        
        |  |     +-- Lampsilis_ornata     
        +--+                              
           |  +----- Venustaconcha        
           +--+                           
              |  +-- Lampsilis_siliquoidea
              +--+                        
                 +-- Lampsilis_powellii   

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

iTtTbiriSoTroTroSiD
220272134--
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

Porto-Hannes et al., 2021 [Publ 4227], fig. 3

iTtTbiriSoTroTroSiD
215011313333
  +-------- Quadrula_quadrula ** 
  |                           
--+-------- Quadrula_verrucosa **
  |                           
  |  +----- Pleurobemini      
  +--+                        
     |  +-- Amblemini         
     +--+                     
        +-- Lampsilini        

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

iTtTbiriSoTroTroSiD
220113773--
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

Jadhav & Jamkhedkar, 2009 [Publ 3697], fig. 11

iTtTbiriSoTroTroSiD
220123113--
1

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

iTtTbiriSoTroTroSiD
220286144--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
22021873--
        +-- Uniomerus **   
  +-----+               
  |     +-- Megalonaias ** 
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini  

Bolotov et al., 2017 [Publ 3764], fig. S1

iTtTbiriSoTroTroSiD
29011237327
  +-- Amblemini         
  |                     
  +-- Ortmanniana       
  |                     
  +-- Lampsilis         
  |                     
  +-- Cambarunio        
  |                     
--+-- Elliptio          
  |                     
  +-- Eurynia           
  |                     
  +-- Pleurobema        
  |                     
  +-- Quadrula_quadrula ** 
  |                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
22017873--
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
220284134--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
214023212332
  +----------- Megalonaias **         
  |                                
  |     +----- Elliptoideus        
  |  +--+                          
--+  |  |  +-- Uniomerus **           
  |  |  +--+                       
  |  |     +-- Pleurobema          
  |  |                             
  |  |  +----- Potamilus_fragilis  
  +--+  |                          
     |  +----- Medionidus          
     |  |                          
     |  +----- Amblema             
     |  |                          
     |  |  +-- Obliquaria          
     +--+--+                       
        |  +-- Plectomerus         
        |                          
        |  +-- Villosa             
        |  |                       
        |  +-- Ellipsaria          
        |  |                       
        +--+-- Lampsilis           
           |                       
           +-- Potamilus_purpuratus
           |                       
           +-- Ptychobranchus      

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

iTtTbiriSoTroTroSiD
220145123--
  +-------- Quadrulina **  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Pleurobemini
     +--+               
        +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220290154--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
220251134--
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
22017473--
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

Inoue et al., 2020 [Publ 3941], fig. 4

iTtTbiriSoTroTroSiD
220139383--
  +----------- Quadrulina **  
--+                        
  |  +-------- Reginaia    
  +--+                     
     |  +----- Pleurobemini
     +--+                  
        |  +-- Amblema     
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
1101611--
-- Quadrula_quadrula **

Campbell et al., 2008 [Publ 2675], fig. 4

iTtTbiriSoTroTroSiD
110124243--
  +----------------------- Uniomerus **                
  |                                                 
  +----------------------- Amblemini                
  |                                                 
  |  +-------------------- Lampsilini               
  |  |                                              
  |  +-------------------- Eurynia                  
  |  |                                              
  |  +-------------------- Fusconaia_cerina         
--+  |                                              
  |  +-------------------- Fusconaia_flava          
  |  |                                              
  |  +-------------------- Hemistena                
  |  |                                              
  |  +-------------------- Plethobasus              
  |  |                                              
  |  +-------------------- Parvaspina               
  |  |                                              
  +--+-------------------- Pleurobema_cordatum      
     |                                              
     +-------------------- Pleuronaia               
     |                                              
     +-------------------- Pleurobema_oviforme__2   
     |                                              
     +-------------------- Pleurobema_stabile       
     |                                              
     +-------------------- Elliptio                 
     |                                              
     |  +----------------- Pleurobema_decisum       
     |  |                                           
     |  +----------------- Pleurobema_sintoxia      
     |  |                                           
     |  +----------------- Pleurobema_clava         
     +--+                                           
        +----------------- Pleurobema_oviforme__1   
        |                                           
        |  +-------------- Pleurobema_beadleianum   
        |  |                                        
        +--+  +----------- Pleurobema_pyriforme     
           |  |                                     
           +--+  +-------- Pleurobema_rubellum      
              |  |                                  
              |  +-------- Pleurobema_hanleyianum__2
              +--+                                  
                 +-------- Pleurobema_georgianum__2 
                 |                                  
                 |  +----- Pleurobema_hanleyianum__1
                 +--+                               
                    |  +-- Pleurobema_georgianum__1 
                    +--+                            
                       +-- Pleurobema_athearni      

Kandl et al., 2001 [Publ 3384], fig. 5

iTtTbiriSoTroTroSiD
1101882--
  +-- Amblemini   
  |               
--+-- Quadrulina **  
  |               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
110129163--
     +-- Quadrulina **  
  +--+               
  |  +-- Pleurobemini
--+                  
  |  +-- Amblemini   
  +--+               
     +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
11013861--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Campbell et al., 2008 [Publ 2675], fig. 5

iTtTbiriSoTroTroSiD
110124243--
  +----------- Uniomerus **                
  |                                     
  +----------- Amblemini                
  |                                     
  |        +-- Elliptio_arctata         
  |  +-----+                            
  |  |     +-- Elliptio_crassidens      
  |  |                                  
  |  |  +----- Pleurobema_decisum       
--+  |  |                               
  |  |  +----- Pleurobema_rubellum      
  |  |  |                               
  |  |  +----- Pleurobema_georgianum__2 
  |  |  |                               
  |  |  +----- Pleurobema_stabile       
  |  |  |                               
  |  |  +----- Pleurobema_pyriforme     
  |  |  |                               
  +--+  +----- Pleurobema_oviforme__2   
     |  |                               
     |  +----- Pleurobema_hanleyianum__2
     |  |                               
     |  +----- Pleuronaia               
     |  |                               
     |  +----- Pleurobema_georgianum__3 
     |  |                               
     |  +----- Pleurobema_cordatum      
     |  |                               
     |  +----- Pleurobema_beadleianum   
     |  |                               
     |  +----- Hemistena                
     +--+                               
        +----- Fusconaia_flava          
        |                               
        +----- Fusconaia_cerina         
        |                               
        +----- Eurynia                  
        |                               
        +----- Elliptio_arca            
        |                               
        +----- Lampsilini               
        |                               
        |  +-- Plethobasus              
        +--+                            
        |  +-- Parvaspina               
        |                               
        |  +-- Pleurobema_sintoxia      
        |  |                            
        +--+-- Pleurobema_oviforme__1   
        |  |                            
        |  +-- Pleurobema_clava         
        |                               
        |  +-- Pleurobema_athearni      
        |  |                            
        +--+-- Pleurobema_georgianum__1 
           |                            
           +-- Pleurobema_hanleyianum__1

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

iTtTbiriSoTroTroSiD
1101511--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11014551--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Wu et al., 2016 [Publ 3711], fig. 1

iTtTbiriSoTroTroSiD
11012021--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
110133103--
  +-------- Toxolasma   
  |                     
  +-------- Amblemini   
  |                     
  |     +-- Glebula     
  +-----+               
--+     +-- Cyrtonaias  
  |                     
  |     +-- Quadrulina **  
  +-----+               
  |     +-- Pleurobemini
  |                     
  |  +----- Potamilus   
  +--+                  
     |  +-- Ortmanniana 
     +--+               
        +-- Ligumia     

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

iTtTbiriSoTroTroSiD
1101932--
  +----- Lampsilini  
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Quadrulina **  

Inoue et al., 2020 [Publ 4128], fig. 3

iTtTbiriSoTroTroSiD
110121213--
  +----------- Quadrulina **  
--+                        
  |  +-------- Reginaia    
  +--+                     
     |  +----- Amblema     
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
11012431--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
110130103--
           +-- Glebula     
  +--------+               
  |        +-- Cyrtonaias  
  |                        
--+  +-------- Toxolasma   
  |  |                     
  |  |  +----- Potamilus   
  |  +--+                  
  +--+  |  +-- Ligumia     
     |  +--+               
     |     +-- Ortmanniana 
     |                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Quadrulina **  
        +--+               
           +-- Pleurobemini

Wang et al., 2018 [Publ 3846], fig. 2

iTtTbiriSoTroTroSiD
11013051--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11014411--
  +-- Lampsilini
--+             
  +-- Quadrulina **

Soroka & Burzynski, 2018 [Publ 3762], fig. 3

iTtTbiriSoTroTroSiD
1101611--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11011821--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Mejia-Trujillo & Smith, 2022 [Publ 4274], fig. 2-M

iTtTbiriSoTroTroSiD
1101551--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11011311--
  +-- Amblemini 
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
110130103--
  +-------- Amblemini   
--+                     
  |  +----- Lampsilini  
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
11012921--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Hauck et al., 2023 [Publ 4333], fig. 9

iTtTbiriSoTroTroSiD
110112123--
  +-------- Quadrulina **  
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
11011411--
  +-- Amblemini 
--+             
  +-- Quadrulina **

Inoue et al., 2013 [Publ 3528], fig. 3

iTtTbiriSoTroTroSiD
110116143--
              +-- Pleurobemini
  +-----------+               
  |           +-- Quadrulina **  
  |                           
--+           +-- Paetulunio  
  |  +--------+               
  |  |        +-- Amblemini   
  +--+                        
     |  +-------- Lampsilis   
     |  |                     
     +--+  +----- Leaunio     
        |  |                  
        +--+----- Obovaria    
           |                  
           |  +-- Villosa     
           +--+               
              +-- Cambarunio  

Sheng et al., 2014 [Publ 3704], fig. 2

iTtTbiriSoTroTroSiD
1101721--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11015283--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11013031--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11011852--
  +----- Lampsilini  
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Quadrulina **  

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

iTtTbiriSoTroTroSiD
11019052--
  +----- Quadrulina **  
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Lampsilini  

Plazzi et al., 2011 [Publ 3676], fig. 5

iTtTbiriSoTroTroSiD
11011742--
  +-- Quadrulina **  
  |               
--+-- Pleurobemini
  |               
  +-- Lampsilini  

Xue et al., 2016 [Publ 3659], fig. 1-M

iTtTbiriSoTroTroSiD
1101111--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1101711--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1101711--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11011931--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11013661--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11011021--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Soroka & Burzynski, 2018 [Publ 3762], fig. 3-M

iTtTbiriSoTroTroSiD
1101611--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11013563--
  +-- Amblemini  
  |              
  +-- Ortmanniana
  |              
  +-- Lampsilis  
  |              
--+-- Cambarunio 
  |              
  +-- Elliptio   
  |              
  +-- Pleurobema 
  |              
  +-- Quadrulina ** 

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

iTtTbiriSoTroTroSiD
11011121--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011121--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11012221--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Wei et al., 2016 [Publ 3665], fig. 0

iTtTbiriSoTroTroSiD
1101511--
-- Quadrula_quadrula **

Smith et al., 2019 [Publ 3951], fig. 2

iTtTbiriSoTroTroSiD
110140403--
  +-------------------------------------- Quadrulina **             
  |                                                              
  |                                   +-- Amblema                
--+  +--------------------------------+                          
  |  |                                +-- Reginaia               
  |  |                                                           
  +--+  +-------------------------------- Plectomerus            
     |  |                                                        
     |  |  +----------------------------- Toxolasma              
     +--+  |                                                     
        |  |                       +----- Popenaiadini           
        |  |  +--------------------+                             
        +--+  |                    |  +-- Glebula                
           |  |                    +--+                          
           |  |                       +-- Cyrtonaias             
           |  |                                                  
           +--+  +----------------------- Obliquaria             
              |  |                                               
              |  |                    +-- Medionidus             
              |  |                 +--+                          
              |  |                 |  +-- Lemiox                 
              +--+  +--------------+                             
                 |  |              |  +-- Cyprogenia             
                 |  |              +--+                          
                 |  |                 +-- Ptychobranchus         
                 |  |                                            
                 +--+           +-------- Potamilus              
                    |  +--------+                                
                    |  |        |  +----- Atlanticoncha          
                    |  |        +--+                             
                    |  |           |  +-- Ellipsaria             
                    |  |           +--+                          
                    +--+              +-- Truncilla              
                       |                                         
                       |              +-- Hamiota                
                       |  +-----------+                          
                       |  |           +-- Villosa                
                       |  |                                      
                       +--+  +----------- Ligumia                
                          |  |                                   
                          |  |     +----- Venustaconcha          
                          +--+  +--+                             
                             |  |  |  +-- Obovaria               
                             |  |  +--+                          
                             +--+     +-- Epioblasma             
                                |                                
                                |  +----- Ortmanniana_pectorosa  
                                +--+                             
                                   |  +-- Ortmanniana_ligamentina
                                   +--+                          
                                      +-- Lampsilis              

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

iTtTbiriSoTroTroSiD
11011621--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Krebs, 2004 [Publ 2692], fig. 2

iTtTbiriSoTroTroSiD
1101422--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012431--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11014411--
  +-- Lampsilini
--+             
  +-- Quadrulina **

Smith et al., 2020 [Publ 4129], fig. 2-3

iTtTbiriSoTroTroSiD
110127274--
  +----------- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
11011021--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11013741--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11014751--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11012121--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11011821--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11013931--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1101611--
-- Quadrula_quadrula **

Smith et al., 2020 [Publ 4129], fig. 1

iTtTbiriSoTroTroSiD
110127274--
  +----------- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
110129163--
     +-- Quadrulina **  
  +--+               
  |  +-- Pleurobemini
--+                  
  |  +-- Amblemini   
  +--+               
     +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
11014031--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Inoue et al., 2020 [Publ 4128], fig. 2

iTtTbiriSoTroTroSiD
110122223--
  +----------- Quadrulina **  
--+                        
  |  +-------- Reginaia    
  +--+                     
     |  +----- Amblema     
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
11011321--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11014351--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Bolotov et al., 2018 [Publ 3826], fig. 2

iTtTbiriSoTroTroSiD
11018263--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
110130103--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011551--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Chen et al., 2020 [Publ 4170], fig. 1

iTtTbiriSoTroTroSiD
11013361--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Xiang et al., 2025 [Publ 4433], fig. 7

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012221--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1101911--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11019052--
  +----- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

Hou et al., 2025 [Publ 4445], fig. 5

iTtTbiriSoTroTroSiD
11015211--
  +-- Lampsilini
--+             
  +-- Quadrulina **

Smith et al., 2020 [Publ 4136], fig. 1

iTtTbiriSoTroTroSiD
110119191--
  +-- Quadrulina **  
--+               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
110112133--
  +-- Amblemini   
  |               
  +-- Pleurobemini
--+               
  +-- Lampsilini  
  |               
  +-- Quadrulina **  

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

iTtTbiriSoTroTroSiD
11013131--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11011921--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Froufe et al., 2015 [Publ 3664], fig. 1-M

iTtTbiriSoTroTroSiD
1101711--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11011531--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11013942--
  +----- Quadrulina **
--+                
  |  +-- Amblemini 
  +--+             
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
110113113--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
110130103--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012121--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11017511--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11014151--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Mulvey et al., 1997 [Publ 3390], fig. 1_3

iTtTbiriSoTroTroSiD
1101431--
  +-- Megalonaias **
--+              
  +-- Amblemini  

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

iTtTbiriSoTroTroSiD
11011933--
-- Quadrula_quadrula **

Burzynski et al., 2017 [Publ 3774], fig. 1-bottom

iTtTbiriSoTroTroSiD
11011211--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Froufe et al., 2015 [Publ 3664], fig. 1

iTtTbiriSoTroTroSiD
11011111--
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
11014851--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Wu et al., 2018 [Publ 4143], fig. 2

iTtTbiriSoTroTroSiD
11012121--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11013043--
  +-------- Quadrulina **  
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
11014351--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11014031--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1101521--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
110168114--
  +----------- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini  

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

iTtTbiriSoTroTroSiD
1101811--
  +-- Lampsilini
--+             
  +-- Quadrulina **

Burzynski et al., 2017 [Publ 3774], fig. 1-top

iTtTbiriSoTroTroSiD
11011211--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11013543--
  +-------- Pleurobemini
--+                     
  |  +----- Quadrulina **  
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini  

Perkins et al., 2017 [Publ 3777], fig. 3

iTtTbiriSoTroTroSiD
110123232--
  +-- Amblemini   
  |               
--+-- Uniomerus **   
  |               
  +-- Pleurobemini

Zhou et al., 2017 [Publ 3788], fig. 2

iTtTbiriSoTroTroSiD
11011521--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Wang et al., 2018 [Publ 3865], fig. 2

iTtTbiriSoTroTroSiD
11013051--
  +-- Quadrulina **
--+             
  +-- Lampsilini

Chen et al., 2012 [Publ 3624], fig. 4

iTtTbiriSoTroTroSiD
1101621--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
110130103--
  +-------- Amblemini   
--+                     
  |  +----- Lampsilini  
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

Kandl et al., 2001 [Publ 3384], fig. 4

iTtTbiriSoTroTroSiD
1101551--
  +-- Elliptio  
  |             
--+-- Uniomerus ** 
  |             
  +-- Pleurobema

Dai et al., 2024 [Publ 4386], fig. 6

iTtTbiriSoTroTroSiD
11014031--
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
11013543--
  +----- Quadrulina **  
  |                  
--+----- Pleurobemini
  |                  
  |  +-- Amblemini   
  +--+               
     +-- Lampsilini  
 
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