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Thu 3 September 2025

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

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

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Bivalvia | Unionida | Unionoidea | Unionidae | Ambleminae | Pleurobemini

genus Hemistena Rafinesque, 1820: cladomics
     1 Recent valid species (1 total species)

Cladomics Summary

number of species = 1 (1 total)
analyzed in 26 cladograms in 13 publications
maximum number of terminals in an analysis = 1 (mean = 1, median = 1)


Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14

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

iTtTbiriSoTroTroSiD
1101630648--
  +-------------------------------------------------------------- Elliptoideus             
  |                                                                                        
  |  +----------------------------------------------------------- Parvaspina               
  |  |                                                                                     
  |  |                                                        +-- Elliptio_fisheriana      
--+  |  +-----------------------------------------------------+                            
  |  |  |                                                     +-- Elliptio_nasutidus       
  |  |  |                                                                                  
  |  |  |  +----------------------------------------------------- Elliptio_spinosa         
  |  |  |  |                                                                               
  +--+  |  |                       +----------------------------- Elliptio_dariensis__5    
     |  |  |                       |                                                       
     |  |  |                       |                       +----- Elliptio_arctata__3      
     |  |  |                       |                       |                               
     |  |  |                       |                       +----- Elliptio_crassidens__4   
     |  |  |                       |  +--------------------+                               
     |  |  |                       |  |                    +----- Elliptio_mcmichaeli__2   
     |  |  |                       |  |                    |                               
     |  |  |                       |  |                    |  +-- Elliptio_crassidens__5   
     +--+  |                       |  |                    +--+                            
        |  |  +--------------------+  |                       +-- Elliptio_roanokensis     
        |  |  |                    |  |                                                    
        |  |  |                    |  |                    +----- Elliptio_complanata__2   
        |  |  |                    |  |                    |                               
        |  |  |                    |  |                    +----- Elliptio_pullata__2      
        |  |  |                    |  |                    |                               
        |  |  |                    |  |                    +----- Elliptio_icterina__3     
        |  |  |                    |  |                    |                               
        |  |  |                    |  +--------------------+  +-- Elliptio_dariensis__4    
        |  |  |                    |  |                    +--+                            
        |  |  |                    |  |                    |  +-- Elliptio_jayensis__2     
        |  |  |                    +--+                    |                               
        |  |  |                       |                    |  +-- Elliptio_congaraea__2    
        +--+  |                       |                    |  |                            
           |  |                       |                    |  +-- Elliptio_crassidens__3   
           |  |                       |                    +--+                            
           |  |                       |                       +-- Elliptio_fumata__2       
           |  |                       |                       |                            
           |  |                       |                       +-- Elliptio_mcmichaeli__1   
           |  |                       |                                                    
           |  |                       |  +----------------------- Elliptio_crassidens__2   
           |  |                       |  |                                                 
           |  |                       |  |                    +-- Elliptio_arca__2         
           |  |                       |  +--------------------+                            
           |  |                       |  |                    +-- Elliptio_crassidens__1   
           |  |                       |  |                                                 
           |  |                       |  |              +-------- Elliptio_icterina__2     
           |  |                       |  |  +-----------+                                  
           |  |                       +--+  |           |  +----- Elliptio_dariensis__3    
           |  |                          |  |           +--+                               
           |  |                          |  |              |  +-- Elliptio_dariensis__2    
           |  |                          |  |              +--+                            
           |  |                          |  |                 +-- Elliptio_mediocris       
           |  |                          |  |                                              
           |  |                          |  |                 +-- Elliptio_folliculata__2  
           |  |                          |  |     +-----------+                            
           |  |                          |  |     |           +-- Elliptio_purpurella__2   
           |  |                          +--+     |                                        
           |  |                             |     |           +-- Elliptio_jayensis__1     
           +--+                             |  +--+-----------+                            
              |                             |  |  |           +-- Elliptio_pullata__1      
              |                             |  |  |                                        
              |                             |  |  |  +----------- Elliptio_arca__1         
              |                             |  |  +--+                                     
              |                             |  |     |  +-------- Elliptio_arctata__2      
              |                             |  |     +--+                                  
              |                             |  |        |  +----- Elliptio_fumata__1       
              |                             |  |        +--+                               
              |                             +--+           |  +-- Elliptio_arctata__1      
              |                                |           +--+                            
              |                                |              +-- Elliptio_purpurella__1   
              |                                |                                           
              |                                |              +-- Elliptio_congaraea__1    
              |                                |              |                            
              |                                |     +--------+-- Elliptio__1              
              |                                |     |        |                            
              |                                |     |        +-- Elliptio__2              
              |                                |     |                                     
              |                                +-----+        +-- Elliptio_complanata__1   
              |                                      |  +-----+                            
              |                                      |  |     +-- Elliptio_hopetonensis__1 
              |                                      |  |                                  
              |                                      +--+  +----- Elliptio_dariensis__1    
              |                                         |  |                               
              |                                         |  +----- Elliptio_producta        
              |                                         +--+                               
              |                                            |  +-- Elliptio_hopetonensis__2 
              |                                            |  |                            
              |                                            +--+-- Elliptio_hopetonensis__3 
              |                                               |                            
              |                                               +-- Elliptio_icterina__1     
              |                                                                            
              |                                            +----- Elliptio_shepardiana     
              |  +-----------------------------------------+                               
              |  |                                         |  +-- Elliptio_angustata       
              |  |                                         +--+                            
              |  |                                            +-- Elliptio_folliculata__1  
              |  |                                                                         
              +--+  +-------------------------------------------- Eurynia                  
                 |  |                                                                      
                 |  |                                         +-- Hemistena_lata **           
                 |  |  +--------------------------------------+                            
                 |  |  |                                      +-- Plethobasus_cyphyus      
                 +--+  |                                                                   
                    |  |  +-------------------------------------- Elliptio_fraterna        
                    |  |  |                                                                
                    |  |  |     +-------------------------------- Pleurobema_plenum        
                    |  |  |     |                                                          
                    +--+  |  +--+  +----------------------------- Pleurobema_stabile       
                       |  |  |  |  |                                                       
                       |  |  |  |  |  +-------------------------- Pleuronaia               
                       |  |  |  +--+  |                                                    
                       |  |  |     |  |                       +-- Pleurobema_decisum       
                       |  |  |     |  |                 +-----+                            
                       |  |  |     |  |                 |     +-- Pleurobema_hanleyianum   
                       |  |  |     +--+  +--------------+                                  
                       +--+  |        |  |              |  +----- Pleurobema_pyriforme     
                          |  |        |  |              +--+                               
                          |  |        |  |                 |  +-- Pleurobema_athearni      
                          |  |        |  |                 +--+                            
                          |  |        |  |                    +-- Pleurobema_georgianum    
                          |  |        +--+                                                 
                          |  |           |                    +-- Pleurobema_clava__1      
                          |  |           |                 +--+                            
                          |  |           |                 |  +-- Pleurobema_oviforme__1   
                          |  |           |  +--------------+                               
                          |  |           |  |              |  +-- Pleurobema_clava__2      
                          |  |           |  |              +--+                            
                          |  |           +--+                 +-- Pleurobema_oviforme__2   
                          +--+              |                                              
                             |              |  +----------------- Pleurobema_beadleianum__2
                             |              +--+                                           
                             |                 |  +-------------- Pleurobema_rubellum      
                             |                 +--+                                        
                             |                    |  +----------- Pleurobema_strodeanum__2 
                             |                    +--+                                     
                             |                       |  +-------- Pleurobema_beadleianum__1
                             |                       +--+                                  
                             |                          |  +----- Pleurobema_taitianum     
                             |                          +--+                               
                             |                             |  +-- Pleurobema_strodeanum__1 
                             |                             +--+                            
                             |                                +-- Pleurobema_perovatum     
                             |                                                             
                             |                             +----- Pleurobema_riddellii     
                             |           +-----------------+                               
                             |           |                 |  +-- Plethobasus_cooperianus  
                             |           |                 +--+                            
                             |           |                    +-- Pleurobema_sintoxia      
                             |           |                                                 
                             +-----------+              +-------- Pleurobema_cordatum      
                                         |  +-----------+                                  
                                         |  |           |  +----- Fusconaia_subrotunda     
                                         |  |           +--+                               
                                         |  |              |  +-- Fusconaia_cor            
                                         |  |              +--+                            
                                         +--+                 +-- Fusconaia_cuneolus       
                                            |                                              
                                            |  +----------------- Fusconaia_ozarkensis     
                                            |  |                                           
                                            |  |           +----- Fusconaia_escambia__2    
                                            |  |  +--------+                               
                                            +--+  |        |  +-- Fusconaia_burkei         
                                               |  |        +--+                            
                                               |  |           +-- Fusconaia_escambia__1    
                                               |  |                                        
                                               +--+        +----- Fusconaia_masoni         
                                                  |        |                               
                                                  |  +-----+  +-- Fusconaia_cerina__1      
                                                  |  |     |  |                            
                                                  |  |     +--+-- Fusconaia_cerina__2      
                                                  |  |        |                            
                                                  +--+        +-- Fusconaia_flava__2       
                                                     |                                     
                                                     |        +-- Fusconaia_askewi         
                                                     |  +-----+                            
                                                     |  |     +-- Fusconaia_chunii         
                                                     +--+                                  
                                                        |  +----- Fusconaia_flava__1       
                                                        +--+                               
                                                           |  +-- Fusconaia_iheringi       
                                                           +--+                            
                                                              +-- Fusconaia_mitchelli      

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

iTtTbiriSoTroTroSiD
110137208--
  +----------------- Elliptio             
  |                                       
  |  +-------------- Parvaspina           
--+  |                                    
  |  +-------------- Plethobasus          
  |  |                                    
  |  |           +-- Hemistena_lata **       
  +--+-----------+                        
     |           +-- Eurynia              
     |                                    
     |  +----------- Fusconaia            
     |  |                                 
     |  |     +----- Pleurobema_riddellii 
     +--+  +--+                           
        |  |  |  +-- Pleurobema_sintoxia  
        |  |  +--+                        
        +--+     +-- Pleurobema_cordatum  
           |                              
           |  +----- Pleuronaia           
           |  |                           
           +--+  +-- Pleurobema_pyriforme 
              |  |                        
              +--+-- Pleurobema_strodeanum
                 |                        
                 +-- Pleurobema_clava     

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

iTtTbiriSoTroTroSiD
11015644--
  +-------- Pleuronaia    
  |                       
--+-------- Pleurobema    
  |                       
  |  +----- Fusconaia     
  +--+                    
     |  +-- Hemistena_lata **
     +--+                 
        +-- Eurynia       

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

iTtTbiriSoTroTroSiD
110141248--
  +-------------------- Elliptoideus        
  |                                         
  |  +----------------- Pleurobema_stabile  
--+  |                                      
  |  |           +----- Pleurobema_pyriforme
  |  |  +--------+                          
  +--+  |        |  +-- Pleurobema_clava    
     |  |        +--+                       
     |  |           +-- Pleurobema_decisum  
     |  |                                   
     +--+           +-- Pleurobema_cordatum 
        |  +--------+                       
        |  |        +-- Pleurobema_sintoxia 
        |  |                                
        |  |     +----- Pleuronaia          
        +--+  +--+                          
           |  |  |  +-- Parvaspina          
           |  |  +--+                       
           |  |     +-- Fusconaia           
           +--+                             
              |     +-- Eurynia             
              |  +--+                       
              |  |  +-- Hemistena_lata **      
              +--+                          
                 |  +-- Plethobasus         
                 +--+                       
                    +-- Elliptio            

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

iTtTbiriSoTroTroSiD
11014075--
  +----------- Parvaspina    
  |                          
--+        +-- Eurynia       
  |  +-----+                 
  |  |     +-- Hemistena_lata **
  +--+                       
     |  +----- Pleurobema    
     +--+                    
        |  +-- Pleuronaia    
        +--+                 
           +-- Fusconaia     

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

iTtTbiriSoTroTroSiD
1101104588--
  +----------------------------------------- Elliptoideus             
  |                                                                   
  |  +-------------------------------------- Parvaspina               
  |  |                                                                
--+  |  +----------------------------------- Plethobasus              
  |  |  |                                                             
  |  |  +----------------------------------- Eurynia                  
  |  |  |                                                             
  |  |  +----------------------------------- Hemistena_lata **           
  +--+  |                                                             
     |  +----------------------------------- Elliptio                 
     |  |                                                             
     |  |  +-------------------------------- Pleuronaia               
     |  |  |                                                          
     |  |  |                             +-- Pleurobema_cordatum      
     |  |  |  +--------------------------+                            
     +--+  |  |                          +-- Fusconaia                
        |  |  |                                                       
        |  |  |  +-------------------------- Pleurobema_plenum        
        |  |  |  |                                                    
        |  |  |  |  +----------------------- Pleurobema_stabile       
        |  |  |  |  |                                                 
        |  |  |  |  |                 +----- Pleurobema_pyriforme     
        |  |  +--+  |              +--+                               
        |  |  |  |  |              |  |  +-- Pleurobema_georgianum__2 
        |  |  |  |  |              |  +--+                            
        |  |  |  |  |  +-----------+     +-- Pleurobema_athearni      
        |  |  |  |  |  |           |                                  
        +--+  |  +--+  |           |  +----- Pleurobema_decisum       
           |  |     |  |           |  |                               
           |  |     |  |           +--+  +-- Pleurobema_georgianum__1 
           |  |     |  |              |  |                            
           |  |     |  |              +--+-- Pleurobema_hanleyianum__1
           |  |     |  |                 |                            
           |  |     |  |                 +-- Pleurobema_hanleyianum__2
           |  |     |  |                                              
           |  |     +--+     +-------------- Pleurobema_beadleianum__2
           |  |        |     |                                        
           |  |        |  +--+  +----------- Pleurobema_strodeanum__2 
           |  |        |  |  |  |                                     
           |  |        |  |  +--+----------- Pleurobema_rubellum      
           |  |        |  |     |                                     
           |  |        |  |     |  +-------- Pleurobema_beadleianum__1
           |  |        |  |     +--+                                  
           +--+        |  |        |  +----- Pleurobema_taitianum     
              |        |  |        +--+                               
              |        |  |           |  +-- Pleurobema_perovatum     
              |        +--+           +--+                            
              |           |              +-- Pleurobema_strodeanum__1 
              |           |                                           
              |           |           +----- Pleurobema_oviforme__6   
              |           |           |                               
              |           |        +--+----- Pleurobema_clava__3      
              |           |        |  |                               
              |           |        |  |  +-- Pleurobema_oviforme__5   
              |           |        |  +--+                            
              |           +--------+     +-- Pleurobema_clava__2      
              |                    |                                  
              |                    |  +----- Pleurobema_oviforme__2   
              |                    |  |                               
              |                    |  +----- Pleurobema_oviforme__3   
              |                    +--+                               
              |                       +----- Pleurobema_oviforme__4   
              |                       |                               
              |                       |  +-- Pleurobema_clava__1      
              |                       +--+                            
              |                          +-- Pleurobema_oviforme__1   
              |                                                       
              |           +----------------- Pleurobema_riddellii     
              |           |                                           
              |           |  +-------------- Pleurobema_sintoxia__34  
              |           |  |                                        
              |           |  |  +----------- Pleurobema_sintoxia__31  
              |           |  |  |                                     
              +-----------+  |  +----------- Pleurobema_sintoxia__32  
                          |  |  |                                     
                          |  |  +----------- Pleurobema_sintoxia__33  
                          |  |  |                                     
                          |  |  |  +-------- Pleurobema__4            
                          |  |  |  |                                  
                          |  |  |  +-------- Pleurobema__5            
                          +--+  |  |                                  
                             |  |  +-------- Pleurobema_sintoxia__2   
                             |  |  |                                  
                             |  |  +-------- Pleurobema_sintoxia__3   
                             |  |  |                                  
                             |  |  +-------- Pleurobema__8            
                             |  |  |                                  
                             |  |  +-------- Pleurobema__9            
                             |  |  |                                  
                             |  |  +-------- Pleurobema__10           
                             |  |  |                                  
                             |  |  +-------- Pleurobema_sintoxia__5   
                             +--+  |                                  
                                |  +-------- Pleurobema_sintoxia__6   
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__7   
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__8   
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__9   
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__10  
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__11  
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__12  
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__13  
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__14  
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__15  
                                |  |                                  
                                |  +-------- Pleurobema_sintoxia__16  
                                +--+                                  
                                   +-------- Pleurobema_sintoxia__17  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__18  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__19  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__20  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__21  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__22  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__23  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__24  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__25  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__26  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__27  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__28  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__29  
                                   |                                  
                                   +-------- Pleurobema_sintoxia__30  
                                   |                                  
                                   |     +-- Pleurobema__6            
                                   +-----+                            
                                   |     +-- Pleurobema_sintoxia__1   
                                   |                                  
                                   |     +-- Pleurobema__7            
                                   +-----+                            
                                   |     +-- Pleurobema_sintoxia__4   
                                   |                                  
                                   |  +----- Pleurobema__3            
                                   +--+                               
                                      |  +-- Pleurobema__1            
                                      +--+                            
                                         +-- Pleurobema__2            

Burlakova et al., 2012 [Publ 3486], fig. 1

iTtTbiriSoTroTroSiD
110140347--
  +----------------------------------- Parvaspina            
  |                                                          
  |                                +-- Plethobasus           
--+  +-----------------------------+                         
  |  |                             +-- Elliptio              
  |  |                                                       
  +--+  +----------------------------- Fusconaia             
     |  |                                                    
     |  |                          +-- Eurynia               
     +--+  +-----------------------+                         
        |  |                       +-- Hemistena_lata **        
        |  |                                                 
        |  |              +----------- Pleuronaia            
        +--+  +-----------+                                  
           |  |           |  +-------- Pleurobema_riddellii  
           |  |           +--+                               
           |  |              |  +----- Pleurobema_sintoxia__2
           |  |              +--+                            
           +--+                 |  +-- Pleurobema_sintoxia__1
              |                 +--+                         
              |                    +-- Pleurobema_cordatum   
              |                                              
              |  +-------------------- Pleurobema_stabile    
              |  |                                           
              +--+  +----------------- Pleurobema_pyriforme  
                 |  |                                        
                 |  |           +----- Pleurobema_georgianum 
                 +--+  +--------+                            
                    |  |        |  +-- Pleurobema_hanleyianum
                    |  |        +--+                         
                    |  |           +-- Pleurobema_decisum    
                    +--+                                     
                       |           +-- Pleurobema_clava      
                       |  +--------+                         
                       |  |        +-- Pleurobema_oviforme   
                       +--+                                  
                          |  +-------- Pleurobema_beadleianum
                          |  |                               
                          +--+     +-- Pleurobema_rubellum   
                             |  +--+                         
                             |  |  +-- Pleurobema_strodeanum 
                             +--+                            
                                |  +-- Pleurobema_perovatum  
                                +--+                         
                                   +-- Pleurobema_taitianum  

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

iTtTbiriSoTroTroSiD
110124217--
-- Hemistena_lata **

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

iTtTbiriSoTroTroSiD
110141248--
  +-------- Elliptoideus  
  |                       
  |  +----- Hemistena_lata **
  |  |                    
--+  +----- Plethobasus   
  |  |                    
  |  +----- Elliptio      
  |  |                    
  +--+  +-- Eurynia       
     +--+                 
     |  +-- Parvaspina    
     |                    
     |  +-- Pleuronaia    
     |  |                 
     +--+-- Pleurobema    
        |                 
        +-- Fusconaia     

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

iTtTbiriSoTroTroSiD
110124217--
-- Hemistena_lata **

Burlakova et al., 2012 [Publ 3486], fig. 2

iTtTbiriSoTroTroSiD
110134347--
  +----------------------------------- Parvaspina            
  |                                                          
  |  +-------------------------------- Pleurobema_plenum     
--+  |                                                       
  |  |                             +-- Plethobasus           
  |  |                          +--+                         
  +--+                          |  +-- Hemistena_lata **        
     |  +-----------------------+                            
     |  |                       |  +-- Eurynia               
     |  |                       +--+                         
     |  |                          +-- Elliptio              
     +--+                                                    
        |  +-------------------------- Pleurobema_cordatum__2
        |  |                                                 
        |  |  +----------------------- Fusconaia             
        +--+  |                                              
           |  |              +-------- Pleurobema_stabile    
           |  |              |                               
           |  |  +-----------+  +----- Pleurobema_riddellii  
           +--+  |           |  |                            
              |  |           +--+  +-- Pleurobema_cordatum__1
              |  |              |  |                         
              |  |              |  +-- Pleurobema_sintoxia__1
              |  |              +--+                         
              +--+                 +-- Pleurobema_sintoxia__2
                 |                 |                         
                 |                 +-- Pleurobema_sintoxia__3
                 |                                           
                 |  +----------------- Pleuronaia            
                 |  |                                        
                 |  |              +-- Pleurobema_georgianum 
                 +--+  +-----------+                         
                    |  |           +-- Pleurobema_pyriforme  
                    |  |                                     
                    +--+           +-- Pleurobema_decisum    
                       |  +--------+                         
                       |  |        +-- Pleurobema_hanleyianum
                       |  |                                  
                       +--+        +-- Pleurobema_clava      
                          |  +-----+                         
                          |  |     +-- Pleurobema_oviforme   
                          |  |                               
                          +--+  +----- Pleurobema_rubellum   
                             |  |                            
                             |  +----- Pleurobema_beadleianum
                             +--+                            
                                +----- Pleurobema_strodeanum 
                                |                            
                                |  +-- Pleurobema_perovatum  
                                +--+                         
                                   +-- Pleurobema_taitianum  

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

iTtTbiriSoTroTroSiD
110123216--
  +----------------- Parvaspina            
  |                                        
  |  +-------------- Elliptio_arca         
  |  |                                     
--+  +-------------- Hemistena_lata **        
  |  |                                     
  |  |           +-- Elliptio_arctata      
  |  +-----------+                         
  |  |           +-- Elliptio_crassidens   
  +--+                                     
     |  +----------- Eurynia               
     |  |                                  
     |  |  +-------- Pleurobema_oviforme   
     |  |  |                               
     |  |  +-------- Pleurobema_clava      
     |  |  |                               
     +--+  +-------- Pleurobema_pyriforme  
        |  |                               
        |  +-------- Pleurobema_georgianum 
        |  |                               
        |  +-------- Pleurobema_beadleianum
        |  |                               
        +--+-------- Pleurobema_hanleyianum
           |                               
           +-------- Pleurobema_decisum    
           |                               
           |     +-- Pleurobema_sintoxia   
           +-----+                         
           |     +-- Pleurobema_rubellum   
           |                               
           |  +----- Elliptio_spinosa      
           |  |                            
           +--+  +-- Fusconaia_escambia    
              |  |                         
              |  +-- Fusconaia_flava       
              +--+                         
                 +-- Fusconaia_cerina      
                 |                         
                 +-- Pleuronaia            

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

iTtTbiriSoTroTroSiD
11012843--
  +----- Hemistena_lata **
--+                    
  |  +-- Elliptio      
  +--+                 
     +-- Fusconaia     

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

iTtTbiriSoTroTroSiD
1101101298--
  +-------------- Elliptoideus  
  |                             
--+  +----------- Hemistena_lata **
  |  |                          
  |  |        +-- Plethobasus   
  +--+  +-----+                 
     |  |     +-- Elliptio      
     |  |                       
     +--+  +----- Pleuronaia    
        |  |                    
        |  +----- Fusconaia     
        +--+                    
           +----- Pleurobema    
           |                    
           |  +-- Eurynia       
           +--+                 
              +-- Parvaspina    

Perkins et al., 2017 [Publ 3777], fig. 2

iTtTbiriSoTroTroSiD
110136346--
  +----------------- Pleurobema    
--+                                
  |  +-------------- Pleuronaia    
  +--+                             
     |  +----------- Fusconaia     
     +--+                          
        |  +-------- Parvaspina    
        +--+                       
           |  +----- Elliptio      
           +--+                    
              |  +-- Eurynia       
              +--+                 
                 +-- Hemistena_lata **

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

iTtTbiriSoTroTroSiD
1101101298--
  +-------------------------------- Elliptoideus             
  |                                                          
  |  +----------------------------- Plethobasus              
--+  |                                                       
  |  +----------------------------- Parvaspina               
  |  |                                                       
  |  +----------------------------- Elliptio                 
  +--+                                                       
     |  +-------------------------- Eurynia                  
     |  |                                                    
     |  +-------------------------- Hemistena_lata **           
     |  |                                                    
     |  |  +----------------------- Fusconaia                
     +--+  |                                                 
        |  |                 +----- Pleurobema_sintoxia__2   
        |  +-----------------+                               
        |  |                 |  +-- Pleurobema_sintoxia__1   
        |  |                 +--+                            
        +--+                    +-- Pleurobema_cordatum      
           |                                                 
           |  +-------------------- Pleuronaia               
           |  |                                              
           |  |           +-------- Pleurobema_rubellum      
           |  |  +--------+                                  
           +--+  |        |  +----- Pleurobema_strodeanum    
              |  |        +--+                               
              |  |           |  +-- Pleurobema_perovatum     
              |  |           +--+                            
              |  |              +-- Pleurobema_taitianum     
              +--+                                           
                 |              +-- Pleurobema_oviforme__1   
                 |              |                            
                 |  +-----------+-- Pleurobema_oviforme__2   
                 |  |           |                            
                 |  |           +-- Pleurobema_clava         
                 +--+                                        
                    |  +----------- Pleurobema_pyriforme     
                    |  |                                     
                    +--+        +-- Pleurobema_athearni      
                       |  +-----+                            
                       |  |     +-- Pleurobema_georgianum__2 
                       +--+                                  
                          |  +----- Pleurobema_decisum       
                          |  |                               
                          +--+  +-- Pleurobema_hanleyianum__1
                             |  |                            
                             +--+-- Pleurobema_hanleyianum__2
                                |                            
                                +-- Pleurobema_georgianum__1 

Campbell et al., 2008 [Publ 2675], fig. 2

iTtTbiriSoTroTroSiD
110125237--
                    +-- Pleurobema_cordatum     
                    |                           
           +--------+-- Pleurobema_sintoxia__1  
           |        |                           
           |        +-- Pleurobema_sintoxia__2  
  +--------+                                    
  |        |  +-------- Fusconaia               
  |        |  |                                 
  |        +--+  +----- Pleuronaia              
  |           |  |                              
  |           |  |  +-- Eurynia                 
  |           +--+  |                           
  |              |  +-- Hemistena_lata **          
  |              |  |                           
--+              +--+-- Plethobasus             
  |                 |                           
  |                 +-- Parvaspina              
  |                 |                           
  |                 +-- Elliptio                
  |                                             
  |  +----------------- Pleurobema_stabile      
  |  |                                          
  |  |  +-------------- Pleurobema_pyriforme    
  |  |  |                                       
  +--+  |           +-- Pleurobema_georgianum__2
     |  +-----------+                           
     |  |           +-- Pleurobema_athearni     
     |  |                                       
     |  |        +----- Pleurobema_decisum      
     +--+--------+                              
        |        |  +-- Pleurobema_hanleyianum  
        |        +--+                           
        |           +-- Pleurobema_georgianum__1
        |                                       
        |           +-- Pleurobema_clava        
        |  +--------+                           
        |  |        +-- Pleurobema_oviforme     
        +--+                                    
           |  +-------- Pleurobema_rubellum     
           |  |                                 
           +--+-------- Pleurobema_strodeanum   
              |                                 
              |  +----- Pleurobema_beadleianum  
              +--+                              
                 |  +-- Pleurobema_taitianum    
                 +--+                           
                    +-- Pleurobema_perovatum    

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

iTtTbiriSoTroTroSiD
110142248--
  +----------------------- Elliptio            
  |                                            
--+  +-------------------- Eurynia             
  |  |                                         
  |  |  +----------------- Parvaspina          
  +--+  |                                      
     |  |              +-- Hemistena_lata **      
     |  +--------------+                       
     +--+              +-- Plethobasus         
        |                                      
        |  +-------------- Pleuronaia          
        |  |                                   
        |  |           +-- Pleurobema_cordatum 
        +--+  +--------+                       
           |  |        +-- Fusconaia           
           |  |                                
           +--+        +-- Pleurobema_stabile  
              |  +-----+                       
              |  |     +-- Pleurobema_sintoxia 
              +--+                             
                 |  +----- Pleurobema_decisum  
                 +--+                          
                    |  +-- Pleurobema_clava    
                    +--+                       
                       +-- Pleurobema_pyriforme

Burlakova et al., 2012 [Publ 3486], fig. 3

iTtTbiriSoTroTroSiD
110135347--
  +----------------------- Hemistena_lata **        
  |                                              
  +----------------------- Pleurobema_stabile    
  |                                              
--+  +-------------------- Eurynia               
  |  |                                           
  |  |           +-------- Pleurobema_riddellii  
  |  |           |                               
  |  |  +--------+  +----- Pleurobema_sintoxia__2
  +--+  |        |  |                            
     |  |        +--+----- Pleurobema_sintoxia__3
     |  |           |                            
     |  |           |  +-- Pleurobema_cordatum   
     |  |           +--+                         
     |  |              +-- Pleurobema_sintoxia__1
     +--+                                        
        |  +-------------- Parvaspina            
        |  |                                     
        |  +-------------- Pleuronaia            
        |  |                                     
        |  +-------------- Fusconaia             
        |  |                                     
        |  |           +-- Plethobasus           
        +--+-----------+                         
           |           +-- Elliptio              
           |                                     
           |           +-- Pleurobema_clava      
           |  +--------+                         
           |  |        +-- Pleurobema_oviforme   
           |  |                                  
           |  |  +-------- Pleurobema_decisum    
           +--+  |                               
              |  +-------- Pleurobema_pyriforme  
              |  |                               
              |  +-------- Pleurobema_hanleyianum
              +--+                               
                 +-------- Pleurobema_georgianum 
                 |                               
                 |  +----- Pleurobema_rubellum   
                 |  |                            
                 |  +----- Pleurobema_beadleianum
                 +--+                            
                    +----- Pleurobema_strodeanum 
                    |                            
                    |  +-- Pleurobema_perovatum  
                    +--+                         
                       +-- Pleurobema_taitianum  

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

iTtTbiriSoTroTroSiD
110119188--
  +-------------- Elliptoideus  
  |                             
--+  +----------- Elliptio      
  |  |                          
  |  +----------- Plethobasus   
  +--+                          
     |  +-------- Parvaspina    
     |  |                       
     |  |     +-- Hemistena_lata **
     +--+-----+                 
        |     +-- Eurynia       
        |                       
        |  +----- Fusconaia     
        +--+                    
           |  +-- Pleuronaia    
           +--+                 
              +-- Pleurobema    

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

iTtTbiriSoTroTroSiD
110141248--
  +-------------------------- Elliptoideus        
  |                                               
--+  +----------------------- Pleurobema_stabile  
  |  |                                            
  +--+  +-------------------- Hemistena_lata **      
     |  |                                         
     +--+  +----------------- Eurynia             
        |  |                                      
        |  |              +-- Pleurobema_cordatum 
        +--+  +-----------+                       
           |  |           +-- Pleurobema_sintoxia 
           |  |                                   
           |  |        +----- Pleurobema_clava    
           +--+  +-----+                          
              |  |     |  +-- Pleurobema_decisum  
              |  |     +--+                       
              |  |        +-- Pleurobema_pyriforme
              +--+                                
                 |        +-- Parvaspina          
                 |  +-----+                       
                 |  |     +-- Pleuronaia          
                 +--+                             
                    |  +----- Fusconaia           
                    +--+                          
                       |  +-- Plethobasus         
                       +--+                       
                          +-- Elliptio            

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

iTtTbiriSoTroTroSiD
11014175--
  +----------- Parvaspina    
  |                          
--+        +-- Eurynia       
  |  +-----+                 
  |  |     +-- Hemistena_lata **
  +--+                       
     |  +----- Pleurobema    
     +--+                    
        |  +-- Pleuronaia    
        +--+                 
           +-- Fusconaia     

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

iTtTbiriSoTroTroSiD
110172408--
  +-------------------------- Elliptoideus          
  |                                                 
  |  +----------------------- Pleurobema_stabile    
  |  |                                              
  |  |                 +----- Pleurobema_riddellii  
--+  |     +-----------+                            
  |  |     |           |  +-- Pleurobema_sintoxia   
  |  |     |           +--+                         
  |  |  +--+              +-- Pleurobema_cordatum   
  |  |  |  |                                        
  +--+  |  |  +-------------- Fusconaia             
     |  |  |  |                                     
     |  |  +--+  +----------- Pleuronaia            
     |  |     |  |                                  
     |  |     +--+        +-- Eurynia               
     |  |        |  +-----+                         
     |  |        |  |     +-- Parvaspina            
     |  |        +--+                               
     +--+           |  +----- Elliptio              
        |           +--+                            
        |              |  +-- Hemistena_lata **        
        |              +--+                         
        |                 +-- Plethobasus           
        |                                           
        |                 +-- Pleurobema_clava      
        |     +-----------+                         
        |     |           +-- Pleurobema_oviforme   
        |     |                                     
        +-----+  +----------- Pleurobema_pyriforme  
              |  |                                  
              |  |        +-- Pleurobema_athearni   
              |  |     +--+                         
              +--+     |  +-- Pleurobema_georgianum 
                 |  +--+                            
                 |  |  |  +-- Pleurobema_decisum    
                 |  |  +--+                         
                 |  |     +-- Pleurobema_hanleyianum
                 +--+                               
                    |  +----- Pleurobema_beadleianum
                    |  |                            
                    |  +----- Pleurobema_strodeanum 
                    +--+                            
                       +----- Pleurobema_rubellum   
                       |                            
                       |  +-- Pleurobema_perovatum  
                       +--+                         
                          +-- Pleurobema_taitianum  

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

iTtTbiriSoTroTroSiD
110172408--
  +----------- Elliptoideus  
  |                          
  |  +-------- Elliptio      
--+  |                       
  |  |     +-- Eurynia       
  |  +-----+                 
  |  |     +-- Parvaspina    
  +--+                       
     |     +-- Hemistena_lata **
     +-----+                 
     |     +-- Plethobasus   
     |                       
     |  +----- Pleuronaia    
     +--+                    
        |  +-- Fusconaia     
        +--+                 
           +-- Pleurobema    

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

iTtTbiriSoTroTroSiD
11012843--
  +----- Hemistena_lata **
--+                    
  |  +-- Elliptio      
  +--+                 
     +-- Fusconaia     

Campbell et al., 2008 [Publ 2675], fig. 6

iTtTbiriSoTroTroSiD
110125237--
  +----------------- Parvaspina    
  |                                
--+  +-------------- Hemistena_lata **
  |  |                             
  +--+           +-- Plethobasus   
     |  +--------+                 
     |  |        +-- Elliptio      
     +--+                          
        |  +-------- Fusconaia     
        +--+                       
           |  +----- Pleurobema    
           +--+                    
              |  +-- Eurynia       
              +--+                 
                 +-- Pleuronaia    
 
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