Bivalvia | Unionida | Unionoidea | Unionidae | Ambleminae

tribe QUADRULINI Ihering, 1901: cladomics

5 genera, 1 subtribe

Cladomics pages are experimental — even more so than the rest of this site!

The purpose of the cladomics pages for each genus and family-group level taxon is to provide statistics and references indicating the extent to which published phylogenetic work supports the classification used here.

The Cladomics Summary provides an overview of the number of studies that have included terminal species representing this taxon as well as an indication of how frequently it has been recovered as monophyletic.

Following the summary, the available published clades representing the taxon are listed.

For more information, visit the Cladomics FARQ (frequently asked rhetorical questions) page. Abbreviations for the various statistics are explained on the FARQ!

Cladomics Summary

number of species = 26 (46 total)
number of subtaxa (rank: subtribe) = 6
analyzed in 196 cladograms in 119 publications
maximum number of terminals in an analysis = 22 (mean = 2.58, 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 of Quadrulini cladogram distances
PCoA Ordination

NJ Tree of Quadrulini cladogram distances
NJ Tree

Cladograms

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

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
22 22 20 3 5 4 4 0 
        +----- Megalonaias ** 
  +-----+                  
  |     |  +-- Uniomerus **   
  |     +--+               
--+        +-- Quadrulina **  
  |                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Lampsilini  
        +--+               
           +-- Popenaiadini

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
11 11 8 3 91 79 4 20 
           +----- Quadrulina **  
  +--------+                  
  |        |  +-- Megalonaias ** 
  |        +--+               
--+           +-- Uniomerus **   
  |                           
  |  +----------- Pleurobemini
  |  |                        
  +--+        +-- Plectomerus 
     |  +-----+               
     |  |     +-- Reginaia    
     +--+                     
        |  +----- Lampsilini  
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Amblema

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

iTtTbiriSoTroTroSiD
19 25 23 3 10 6 3 27 
  +----------- Megalonaias ** 
  |                        
--+        +-- Uniomerus **   
  |  +-----+               
  |  |     +-- Quadrulina **  
  +--+                     
     |  +----- Pleurobemini
     +--+                  
        |  +-- Lampsilini  
        +--+               
           +-- Amblemini

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

iTtTbiriSoTroTroSiD
18 18 15 3 10 6 3 23 
  +-------------- Amblemini   
--+                           
  |  +----------- Lampsilini  
  +--+                        
     |  +-------- Pleurobemini
     +--+                     
        |  +----- Quadrulina **  
        +--+                  
           |  +-- Megalonaias ** 
           +--+               
              +-- Uniomerus **

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

iTtTbiriSoTroTroSiD
16 16 14 3 10 6 3 22 
     +----- Amblemini   
  +--+                  
  |  |  +-- Pleurobemini
  |  +--+               
--+     +-- Lampsilini  
  |                     
  |  +----- Megalonaias ** 
  +--+                  
     |  +-- Uniomerus **   
     +--+               
        +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
14 14 12 3 2 2 2 20 
  +----------- Amblemini   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Megalonaias ** 
     +--+                  
        |  +-- Uniomerus **   
        +--+               
           +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
14 14 11 2 15 15 3 17 
  +----------- Lampsilis   
  |                        
  |  +-------- Obliquaria  
  |  |                     
  |  +-------- Amblema     
  |  |                     
--+  +-------- Reginaia    
  |  |                     
  |  +-------- Potamilus   
  |  |                     
  |  |     +-- Elliptoideus
  |  +-----+               
  |  |     +-- Plectomerus 
  +--+                     
     |     +-- Megalonaias ** 
     +-----+               
     |     +-- Quadrulina **  
     |                     
     |  +----- Medionidus  
     +--+                  
     |  |  +-- Dromus      
     |  +--+               
     |     +-- Cyprogenia  
     |                     
     |  +----- Hemistena   
     +--+                  
        |  +-- Elliptio    
        +--+               
           +-- Fusconaia

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

iTtTbiriSoTroTroSiD
14 14 12 2 0 0 0 20 
  +----- Megalonaias **
  |                 
--+----- Theliderma ** 
  |                 
  |  +-- Quadrula **   
  +--+              
     +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
14 14 12 2 15 15 3 22 
  +-------------------------- Lampsilis   
  |                                       
--+  +----------------------- Potamilus   
  |  |                                    
  |  |                 +----- Medionidus  
  +--+  +--------------+                  
     |  |              |  +-- Dromus      
     |  |              +--+               
     +--+                 +-- Cyprogenia  
        |                                 
        |  +----------------- Obliquaria  
        |  |                              
        +--+  +-------------- Amblema     
           |  |                           
           |  |        +----- Reginaia    
           +--+  +-----+                  
              |  |     |  +-- Elliptoideus
              |  |     +--+               
              |  |        +-- Plectomerus 
              +--+                        
                 |        +-- Megalonaias ** 
                 |  +-----+               
                 |  |     +-- Quadrulina **  
                 +--+                     
                    |  +----- Hemistena   
                    +--+                  
                       |  +-- Elliptio    
                       +--+               
                          +-- Fusconaia

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

iTtTbiriSoTroTroSiD
11 11 8 2 62 62 4 22 
                                   +-- 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
11 11 8 3 91 79 4 20 
                       +----- 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
11 11 9 2 62 62 4 21 
              +-- Uniomerus **   
  +-----------+               
  |           +-- Quadrulina **  
--+                           
  |  +----------- Pleurobemini
  |  |                        
  +--+        +-- Plectomerus 
     |  +-----+               
     |  |     +-- Reginaia    
     +--+                     
        |  +----- Lampsilini  
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Amblema

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

iTtTbiriSoTroTroSiD
7 7 5 2 28 22 3 23 
  +----------- Reginaia    
  |                        
--+        +-- Amblema     
  |  +-----+               
  |  |     +-- Lampsilini  
  +--+                     
     |  +----- Pleurobemini
     +--+                  
        |  +-- Megalonaias ** 
        +--+               
           +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
7 7 2 2 97 75 4 19 
  +-------- Reginaia           
  |                            
  +-------- Amblema            
  |                            
  |  +----- Pleurobemini       
  +--+                         
  |  |  +-- Uniomerus **          
  |  +--+                      
--+     +-- Quadrulina **         
  |                            
  |  +----- Popenaiadini       
  |  |                         
  |  |  +-- Dromus             
  |  |  |                      
  |  |  +-- Epioblasma         
  |  |  |                      
  +--+  +-- Glebula            
     |  |                      
     |  +-- Lampsilis_ovata    
     |  |                      
     |  +-- Lampsilis_virescens
     |  |                      
     |  +-- Lemiox             
     +--+                      
        +-- Obliquaria         
        |                      
        +-- Obovaria_olivaria  
        |                      
        +-- Obovaria_unicolor  
        |                      
        +-- Plectomerus        
        |                      
        +-- Potamilus_alatus   
        |                      
        +-- Potamilus_capax

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

iTtTbiriSoTroTroSiD
7 7 5 2 36 36 4 21 
                                +-- 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

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

iTtTbiriSoTroTroSiD
7 7 5 1 11 0 0 23 
1

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

iTtTbiriSoTroTroSiD
7 29 25 2 31 22 3 41 
  +-------------------------------- Quadrulina **           
  |                                                      
--+  +----------------------------- Megalonaias **          
  |  |                                                   
  +--+  +-------------------------- Toxolasma            
     |  |                                                
     +--+  +----------------------- Amblema              
        |  |                                             
        |  |  +-------------------- Pleurobemini         
        +--+  |                                          
           |  |                 +-- Reginaia             
           |  +-----------------+                        
           +--+                 +-- Obliquaria           
              |                                          
              |  +----------------- Potamilus            
              |  |                                       
              |  |              +-- Ellipsaria           
              +--+  +-----------+                        
                 |  |           +-- Truncilla            
                 |  |                                    
                 +--+  +----------- Lampsilis_cardium    
                    |  |                                 
                    |  |     +----- Ligumia              
                    |  +-----+                           
                    +--+     |  +-- Obovaria             
                       |     +--+                        
                       |        +-- Epioblasma           
                       |                                 
                       |  +-------- Lampsilis_teres      
                       +--+                              
                          |  +----- Lampsilis_siliquoidea
                          +--+                           
                             |  +-- Ortmanniana          
                             +--+                        
                                +-- Lampsilis_higginsii

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

iTtTbiriSoTroTroSiD
6 6 4 1 36 36 4 22 
  +----------------- Quadrulina **  
  |                              
--+  +-------------- Pleurobemini
  |  |                           
  |  |           +-- Popenaiadini
  +--+  +--------+               
     |  |        +-- Amblema     
     |  |                        
     +--+  +-------- Plectomerus 
        |  |                     
        |  |     +-- Cyrtonaias  
        +--+-----+               
           |     +-- Reginaia    
           |                     
           |  +----- Lemiox      
           +--+                  
              |  +-- Lampsilis   
              +--+               
                 +-- Obovaria

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

iTtTbiriSoTroTroSiD
6 6 4 1 36 36 4 22 
  +-------------------- Quadrulina **  
  |                                 
--+  +----------------- Pleurobemini
  |  |                              
  +--+              +-- Reginaia    
     |  +-----------+               
     |  |           +-- Amblema     
     +--+                           
        |  +----------- Popenaiadini
        |  |                        
        +--+  +-------- Cyrtonaias  
           |  |                     
           +--+-------- Plectomerus 
              |                     
              |  +----- Lemiox      
              +--+                  
                 |  +-- Lampsilis   
                 +--+               
                    +-- Obovaria

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

iTtTbiriSoTroTroSiD
6 6 4 2 32 31 3 22 
  +----------------------------- 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
6 6 4 1 36 36 4 22 
  +-------------------- Quadrulina **  
  |                                 
--+  +----------------- Pleurobemini
  |  |                              
  |  |              +-- Popenaiadini
  +--+  +-----------+               
     |  |           +-- Amblema     
     |  |                           
     +--+           +-- Cyrtonaias  
        |  +--------+               
        |  |        +-- Reginaia    
        +--+                        
           |  +-------- Plectomerus 
           +--+                     
              |  +----- Lemiox      
              +--+                  
                 |  +-- Lampsilis   
                 +--+               
                    +-- Obovaria

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

iTtTbiriSoTroTroSiD
5 5 3 3 26 25 4 19 
        +----- Megalonaias ** 
  +-----+                  
  |     |  +-- Uniomerus **   
  |     +--+               
--+        +-- Quadrulina **  
  |                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
4 18 14 3 23 14 3 34 
                          +-- Megalonaias **         
  +-----------------------+                       
  |                       +-- Quadrulina **          
  |                                               
  |                    +----- Elliptoideus        
--+                    |                          
  |  +-----------------+  +-- Eurynia             
  |  |                 |  |                       
  |  |                 +--+-- Pleurobema          
  |  |                    |                       
  +--+                    +-- Uniomerus **           
     |                                            
     |  +-------------------- Amblema             
     |  |                                         
     +--+                 +-- Plectomerus         
        |  +--------------+                       
        |  |              +-- Obliquaria          
        +--+                                      
           |  +-------------- Potamilus_fragilis  
           |  |                                   
           +--+  +----------- Ptychobranchus      
              |  |                                
              +--+  +-------- Medionidus          
                 |  |                             
                 +--+  +----- Potamilus_purpuratus
                    |  |                          
                    +--+----- Lampsilis           
                       |                          
                       |  +-- Ellipsaria          
                       +--+                       
                          +-- Villosa

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

iTtTbiriSoTroTroSiD
4 17 0 3 18 8 3 33 
  +-- Gonideini          
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias **        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus **          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Pleurobema         
  |                      
  +-- Unionini

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

iTtTbiriSoTroTroSiD
4 17 0 3 18 7 3 33 
  +-- Gonideini          
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias **        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus **          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Aculamprotulina    
  |                      
  +-- Unionina

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

iTtTbiriSoTroTroSiD
4 32 31 2 33 14 3 49 
  +-------------- Quadrulina **     
  |                              
  |        +----- Lamprotulini   
--+  +-----+                     
  |  |     |  +-- Chamberlainiini
  |  |     +--+                  
  +--+        +-- Anodontini     
     |                           
     |  +-------- Megalonaias **    
     +--+                        
        |  +----- Pleurobemini   
        +--+                     
           |  +-- Amblemini      
           +--+                  
              +-- Lampsilini

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

iTtTbiriSoTroTroSiD
4 21 20 3 29 17 3 38 
                          +-- 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

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

iTtTbiriSoTroTroSiD
4 4 2 3 195 21 3 20 
                                +----- Quadrulina **        
  +-----------------------------+                        
  |                             |  +-- Megalonaias **       
  |                             +--+                     
--+                                +-- Uniomerus **         
  |                                                      
  |  +-------------------------------- Pleurobemini      
  |  |                                                   
  +--+  +----------------------------- Amblema           
     |  |                                                
     +--+  +-------------------------- Toxolasma         
        |  |                                             
        +--+                       +-- Obliquaria        
           |  +--------------------+                     
           |  |                    +-- Plectomerus       
           +--+                                          
              |  +-------------------- Potamilus_leptodon
              |  |                                       
              +--+  +----------------- Potamilus_alatus  
                 |  |                                    
                 |  |              +-- Medionidus        
                 +--+  +-----------+                     
                    |  |           +-- Ptychobranchus    
                    |  |                                 
                    +--+           +-- Ellipsaria        
                       |  +--------+                     
                       |  |        +-- Truncilla         
                       +--+                              
                          |  +-------- Leaunio           
                          |  |                           
                          +--+     +-- Venustaconcha     
                             |  +--+                     
                             |  |  +-- Cambarunio        
                             +--+                        
                                |  +-- Ortmanniana       
                                +--+                     
                                   +-- Lampsilis

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

iTtTbiriSoTroTroSiD
4 4 2 3 197 21 3 20 
                       +----- Quadrulina **        
  +--------------------+                        
  |                    |  +-- Megalonaias **       
  |                    +--+                     
--+                       +-- Uniomerus **         
  |                                             
  |  +----------------------- Pleurobemini      
  |  |                                          
  +--+  +-------------------- Amblema           
     |  |                                       
     |  |              +----- Toxolasma         
     +--+  +-----------+                        
        |  |           |  +-- Obliquaria        
        |  |           +--+                     
        |  |              +-- Plectomerus       
        +--+                                    
           |  +-------------- Potamilus_leptodon
           |  |                                 
           |  |  +----------- Potamilus_alatus  
           +--+  |                              
              |  |  +-------- Venustaconcha     
              |  |  |                           
              +--+  +-------- Leaunio           
                 |  |                           
                 |  |     +-- Ellipsaria        
                 |  +-----+                     
                 +--+     +-- Truncilla         
                    |                           
                    |     +-- Medionidus        
                    +-----+                     
                    |     +-- Ptychobranchus    
                    |                           
                    |  +----- Cambarunio        
                    +--+                        
                       |  +-- Ortmanniana       
                       +--+                     
                          +-- Lampsilis

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

iTtTbiriSoTroTroSiD
4 10 0 3 15 7 3 26 
  +-- Pyganodon_cataracta
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Lampsilis_straminea
--+                      
  +-- Megalonaias **        
  |                      
  +-- Obliquaria         
  |                      
  +-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus **

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

iTtTbiriSoTroTroSiD
4 4 2 3 0 0 0 22 
  +----- Quadrulina ** 
--+                 
  |  +-- Uniomerus **  
  +--+              
     +-- Megalonaias **

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

iTtTbiriSoTroTroSiD
4 128 73 3 197 21 3 145 
  +----------- Chamberlainia        
  |                                 
  +----------- Megalonaias **          
  |                                 
  +----------- Uniomerus **            
  |                                 
  +----------- Quadrulina **           
  |                                 
  +----------- Sinohyriopsis        
  |                                 
  +----------- Lamprotulini         
  |                                 
  +----------- Gonideini            
  |                                 
  +----------- Pseudodontini        
  |                                 
  +----------- Rectidentini         
  |                                 
  |  +-------- 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        
     |                              
     +-------- Unionina             
     |                              
     +-------- Aculamprotulina      
     |                              
     |     +-- Pletholophus         
     +-----+                        
     |     +-- Sinanodonta          
     |                              
     |  +----- Cuneopsis_celtiformis
     |  |                           
     |  +----- Tchangsinaia         
     |  |                           
     +--+----- Cuneopsis_rufescens  
        |                           
        +----- Nodularia            
        |                           
        |  +-- Cuneopsis_szechenyii 
        +--+                        
           +-- Cuneopsis_heudei

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

iTtTbiriSoTroTroSiD
4 18 16 3 24 14 3 34 
              +-- Megalonaias ** 
  +-----------+               
  |           +-- Quadrulina **  
  |                           
--+        +----- Amblema     
  |  +-----+                  
  |  |     |  +-- Plectomerus 
  |  |     +--+               
  +--+        +-- Lampsilini  
     |                        
     |  +-------- Elliptoideus
     +--+                     
        |  +----- Uniomerus **   
        +--+                  
           |  +-- Eurynia     
           +--+               
              +-- Pleurobema

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

iTtTbiriSoTroTroSiD
3 3 0 1 11 9 3 20 
  +-------- Amblemini   
--+                     
  |  +----- Lampsilini  
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
3 3 1 1 11 9 3 19 
1

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

iTtTbiriSoTroTroSiD
3 3 1 2 35 14 3 19 
  +----- Pleurobemini
--+                  
  |  +-- Megalonaias ** 
  +--+               
     +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
3 3 0 1 11 9 3 20 
  +-------- Lampsilini  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
3 3 1 1 11 9 3 19 
  +-- Quadrulina **  
--+               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 10 4 -- 
     +----- Amblemini   
  +--+                  
  |  |  +-- Pleurobemini
  |  +--+               
  |     +-- Quadrulina **  
--+                     
  |  +----- Obliquaria  
  |  |                  
  |  +----- Toxolasma   
  +--+                  
     +----- Popenaiadini
     |                  
     |  +-- Glebula     
     +--+               
        +-- Cyrtonaias

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

iTtTbiriSoTroTroSiD
2 2 0 1 39 38 3 -- 
  +----------- Quadrulina **  
--+                        
  |  +-------- Reginaia    
  +--+                     
     |  +----- Pleurobemini
     +--+                  
        |  +-- Amblema     
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 54 8 2 -- 
     +-- Quadrulina **  
  +--+               
  |  +-- Uniomerus **   
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 78 7 3 -- 
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
2 2 0 1 45 12 3 -- 
              +-------- Amblemini     
  +-----------+                       
  |           |  +----- Pleurobemini  
  |           +--+                    
  |              |  +-- Obliquaria    
--+              +--+                 
  |                 +-- Quadrulina **    
  |                                   
  |  +----------------- Ptychobranchus
  +--+                                
     |  +-------------- Truncilla     
     +--+                             
        |  +----------- Ortmanniana   
        +--+                          
           |  +-------- Lampsilis     
           +--+                       
              |  +----- Cambarunio    
              +--+                    
                 |  +-- Ligumia       
                 +--+                 
                    +-- Epioblasma

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

iTtTbiriSoTroTroSiD
2 14 0 2 32 12 3 32 
  +----------- Megalonaias **         
  |                                
  |     +----- Elliptoideus        
  |  +--+                          
--+  |  |  +-- Uniomerus **           
  |  |  +--+                       
  |  |     +-- Pleurobema          
  |  |                             
  |  |  +----- Potamilus_fragilis  
  +--+  |                          
     |  +----- Medionidus          
     |  |                          
     |  +----- Amblema             
     |  |                          
     |  |  +-- Obliquaria          
     +--+--+                       
        |  +-- Plectomerus         
        |                          
        |  +-- Villosa             
        |  |                       
        |  +-- Ellipsaria          
        |  |                       
        +--+-- Lampsilis           
           |                       
           +-- Potamilus_purpuratus
           |                       
           +-- Ptychobranchus

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

iTtTbiriSoTroTroSiD
2 2 0 1 45 12 3 -- 
  +-------- Quadrulina **  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Pleurobemini
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 2 1 1 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 94 15 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 44 11 3 -- 
           +-- Obliquaria    
  +--------+                 
  |        +-- Quadrulina **    
  |                          
--+  +-------- Amblemini     
  |  |                       
  |  +-------- Pleurobemini  
  +--+                       
     |  +----- Truncilla     
     |  |                    
     |  +----- Ptychobranchus
     +--+                    
        |  +-- Ortmanniana   
        |  |                 
        |  +-- Cambarunio    
        +--+                 
           +-- Ligumia       
           |                 
           +-- Epioblasma

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 10 4 -- 
        +-- Popenaiadini
  +-----+               
  |     +-- Lampsilini  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Pleurobemini
     +--+               
        +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
2 2 0 2 17 12 4 -- 
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 40 39 3 -- 
  +-------- Quadrulina **  
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 19 18 4 -- 
  +-------------- Quadrulina **  
--+                           
  |  +----------- Pleurobemini
  +--+                        
     |  +-------- Amblema     
     +--+                     
        |  +----- Plectomerus 
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 11 0 1 9 9 3 29 
  +-------- Quadrula **    
  |                     
--+-------- Pustulosa **   
  |                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 19 18 4 -- 
  +-------------- Quadrulina **  
--+                           
  |  +----------- Pleurobemini
  +--+                        
     |  +-------- Amblema     
     +--+                     
        |  +----- Plectomerus 
        +--+                  
           |  +-- Popenaiadini
           +--+               
              +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 72 13 4 -- 
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 9 9 4 -- 
        +-- Popenaiadini
  +-----+               
  |     +-- Lampsilini  
--+                     
  |  +----- Amblemini   
  +--+                  
     |  +-- Pleurobemini
     +--+               
        +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
2 2 0 1 9 7 2 -- 
  +----------------- Glebula            
  |                                     
--+              +-- Amblema            
  |  +-----------+                      
  |  |           +-- Quadrulina **         
  +--+                                  
     |  +----------- Plectomerus        
     +--+                               
        |  +-------- Leaunio            
        +--+                            
           |  +----- Lampsilis_straminea
           +--+                         
              |  +-- Lampsilis_teres    
              +--+                      
                 +-- Potamilus

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

iTtTbiriSoTroTroSiD
2 2 0 1 14 4 2 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
2 2 0 1 18 17 1 -- 
  +-- Quadrulina **  
--+               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
2 2 0 1 74 7 3 -- 
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
2 14 0 2 32 12 3 32 
  +----------------------------- Megalonaias **         
  |                                                  
  |                       +----- Elliptoideus        
--+  +--------------------+                          
  |  |                    |  +-- Uniomerus **           
  |  |                    +--+                       
  +--+                       +-- Pleurobema          
     |                                               
     |  +----------------------- Amblema             
     |  |                                            
     +--+                    +-- Obliquaria          
        |  +-----------------+                       
        |  |                 +-- Plectomerus         
        +--+                                         
           |  +----------------- Medionidus          
           +--+                                      
              |  +-------------- Potamilus_fragilis  
              +--+                                   
                 |  +----------- Ptychobranchus      
                 +--+                                
                    |  +-------- Potamilus_purpuratus
                    +--+                             
                       |  +----- Lampsilis           
                       +--+                          
                          |  +-- Villosa             
                          +--+                       
                             +-- Ellipsaria

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

iTtTbiriSoTroTroSiD
2 2 0 2 51 8 3 -- 
        +-- Quadrulina **  
  +-----+               
  |     +-- Uniomerus **   
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Popenaiadini
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 15 0 1 13 13 3 33 
  +-------- Quadrula_quadrula ** 
  |                           
--+-------- Quadrula_verrucosa **
  |                           
  |  +----- Pleurobemini      
  +--+                        
     |  +-- Amblemini         
     +--+                     
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 72 13 4 -- 
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 90 15 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 86 14 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 137 7 3 -- 
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
2 2 0 2 18 7 3 -- 
        +-- Uniomerus **   
  +-----+               
  |     +-- Megalonaias ** 
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 13 13 4 -- 
     +-------------- Toxolasma            
  +--+                                    
  |  |  +----------- Popenaiadini         
  |  +--+                                 
  |     |  +-------- Amblemini            
  |     +--+                              
  |        |  +----- Pleurobemini         
--+        +--+                           
  |           |  +-- Quadrulina **           
  |           +--+                        
  |              +-- Uniomerus **            
  |                                       
  |     +----------- Potamilus            
  |     |                                 
  +-----+        +-- Lampsilis_cardium    
        |  +-----+                        
        |  |     +-- Lampsilis_ornata     
        +--+                              
           |  +----- Venustaconcha        
           +--+                           
              |  +-- Lampsilis_siliquoidea
              +--+                        
                 +-- Lampsilis_powellii

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

iTtTbiriSoTroTroSiD
2 2 0 2 52 9 2 -- 
     +-- Quadrulina **  
  +--+               
  |  +-- Uniomerus **   
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 10 7 2 -- 
  +-------- Quadrulina ** 
--+                    
  |  +----- Plectomerus
  +--+                 
     |  +-- Amblema    
     +--+              
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 90 15 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 31 31 4 -- 
                          +-- 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

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

iTtTbiriSoTroTroSiD
2 2 0 2 55 13 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 94 15 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 55 13 4 -- 
        +-- Quadrulina **  
  +-----+               
  |     +-- Uniomerus **   
  |                     
--+     +-- Pleurobemini
  |  +--+               
  |  |  +-- Amblemini   
  +--+                  
     |  +-- Popenaiadini
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 84 13 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 74 7 3 -- 
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
2 2 0 1 137 7 3 -- 
     +-- Amblemini   
  +--+               
  |  +-- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
2 2 0 1 23 11 3 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 1 2 1 1 -- 
1

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

iTtTbiriSoTroTroSiD
2 2 0 2 51 10 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 1 44 11 2 -- 
                 +----- Pleurobemini  
  +--------------+                    
  |              |  +-- Obliquaria    
  |              +--+                 
--+                 +-- Quadrulina **    
  |                                   
  |  +----------------- Ptychobranchus
  +--+                                
     |  +-------------- Truncilla     
     +--+                             
        |  +----------- Ortmanniana   
        +--+                          
           |  +-------- Lampsilis     
           +--+                       
              |  +----- Cambarunio    
              +--+                    
                 |  +-- Ligumia       
                 +--+                 
                    +-- Epioblasma

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

iTtTbiriSoTroTroSiD
2 2 0 1 2 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
2 9 0 1 123 7 3 27 
  +-- Amblemini         
  |                     
  +-- Ortmanniana       
  |                     
  +-- Lampsilis         
  |                     
  +-- Cambarunio        
  |                     
--+-- Elliptio          
  |                     
  +-- Eurynia           
  |                     
  +-- Pleurobema        
  |                     
  +-- Quadrula_quadrula ** 
  |                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 72 13 4 -- 
           +-- Uniomerus **   
  +--------+               
  |        +-- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
2 2 0 2 51 13 4 -- 
           +-- Quadrulina **  
  +--------+               
  |        +-- Uniomerus **   
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 43 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 1 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 -- 
  +-- Elliptio  
  |             
--+-- Uniomerus ** 
  |             
  +-- Pleurobema

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 8 8 2 -- 
  +-- Amblemini   
  |               
--+-- Quadrulina **  
  |               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 90 5 2 -- 
  +----- Quadrulina **  
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 24 3 -- 
  +----------- 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

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 4 3 -- 
  +-------- Quadrulina **  
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 14 3 -- 
              +-- Pleurobemini
  +-----------+               
  |           +-- Quadrulina **  
  |                           
--+           +-- Paetulunio  
  |  +--------+               
  |  |        +-- Amblemini   
  +--+                        
     |  +-------- Lampsilis   
     |  |                     
     +--+  +----- Leaunio     
        |  |                  
        +--+----- Obovaria    
           |                  
           |  +-- Villosa     
           +--+               
              +-- Cambarunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 4 3 -- 
  +-------- Pleurobemini
--+                     
  |  +----- Quadrulina **  
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 27 27 4 -- 
  +----------- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 90 5 2 -- 
  +----- Lampsilini  
--+                  
  |  +-- Quadrulina **  
  +--+               
     +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 10 3 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 10 3 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 15 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 10 3 -- 
  +-------- Toxolasma   
  |                     
  +-------- Amblemini   
  |                     
  |     +-- Glebula     
  +-----+               
--+     +-- Cyrtonaias  
  |                     
  |     +-- Quadrulina **  
  +-----+               
  |     +-- Pleurobemini
  |                     
  |  +----- Potamilus   
  +--+                  
     |  +-- Ortmanniana 
     +--+               
        +-- Ligumia

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 1 1 -- 
  +-- Amblemini 
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 3 2 -- 
  +----- Lampsilini  
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 1 1 -- 
  +-- Amblemini 
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 10 3 -- 
           +-- Glebula     
  +--------+               
  |        +-- Cyrtonaias  
  |                        
--+  +-------- Toxolasma   
  |  |                     
  |  |  +----- Potamilus   
  |  +--+                  
  +--+  |  +-- Ligumia     
     |  +--+               
     |     +-- Ortmanniana 
     |                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Quadrulina **  
        +--+               
           +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 2 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 12 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 5 2 -- 
  +----- Lampsilini  
--+                  
  |  +-- Pleurobemini
  +--+               
     +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 17 4 2 -- 
  +-- Quadrulina **  
  |               
--+-- Pleurobemini
  |               
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 37 4 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 16 3 -- 
     +-- Quadrulina **  
  +--+               
  |  +-- Pleurobemini
--+                  
  |  +-- Amblemini   
  +--+               
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 12 12 3 -- 
  +-------- Quadrulina **  
--+                     
  |  +----- Pleurobemini
  +--+                  
     |  +-- Amblemini   
     +--+               
        +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 36 6 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 2 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 8 3 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 8 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 22 3 -- 
  +----------- Quadrulina **  
--+                        
  |  +-------- Reginaia    
  +--+                     
     |  +----- Amblema     
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 15 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 82 6 3 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 6 3 -- 
  +-- Amblemini  
  |              
  +-- Ortmanniana
  |              
  +-- Lampsilis  
  |              
--+-- Cambarunio 
  |              
  +-- Elliptio   
  |              
  +-- Pleurobema 
  |              
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 1 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 1 1 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 75 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 40 3 -- 
  +-------------------------------------- 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

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 12 1 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 4 3 1 -- 
  +-- Megalonaias **
--+              
  +-- Amblemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

Krebs, 2004 [Publ 2692], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 4 2 2 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 16 3 -- 
     +-- Quadrulina **  
  +--+               
  |  +-- Pleurobemini
--+                  
  |  +-- Amblemini   
  +--+               
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 15 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 4 3 -- 
  +----- Quadrulina **  
  |                  
--+----- Pleurobemini
  |                  
  |  +-- Amblemini   
  +--+               
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 10 3 -- 
  +-------- Amblemini   
--+                     
  |  +----- Lampsilini  
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 39 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 10 3 -- 
  +-------- Amblemini   
--+                     
  |  +----- Lampsilini  
  +--+                  
     |  +-- Quadrulina **  
     +--+               
        +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 27 27 4 -- 
  +----------- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 68 11 4 -- 
  +----------- Quadrulina **  
--+                        
  |  +-------- Pleurobemini
  +--+                     
     |  +----- Amblemini   
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 11 3 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 121 3 3 -- 
  +-- Amblemini   
  |               
  +-- Pleurobemini
--+               
  +-- Lampsilini  
  |               
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 48 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 1 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 43 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 24 3 -- 
  +----------------------- 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

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 6 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 38 6 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 23 23 2 -- 
  +-- Amblemini   
  |               
--+-- Uniomerus **   
  |               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 1 1 -- 
  +-- Lampsilini
--+             
  +-- Quadrulina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 19 1 -- 
  +-- Quadrulina **  
--+               
  +-- Pleurobemini

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 47 5 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 2 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 31 3 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 12 1 1 -- 
  +-- Quadrulina **
--+             
  +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 21 3 -- 
  +----------- Quadrulina **  
--+                        
  |  +-------- Reginaia    
  +--+                     
     |  +----- Amblema     
     +--+                  
        |  +-- Popenaiadini
        +--+               
           +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 39 4 2 -- 
  +----- Quadrulina **
--+                
  |  +-- Amblemini 
  +--+             
     +-- Lampsilini

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 3 3 -- 
-- Quadrula_quadrula **