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Page last updated
Wed 1 October 2025

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

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

Search the database pages:

Bivalvia | Unionoida | Unionoidea | Unionidae | Ambleminae | Quadrulini

subtribe QUADRULINA Ihering, 1901: cladomics
     4 genera

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 = 20 (25 total)
number of subtaxa (rank: genus) = 4
analyzed in 185 cladograms in 112 publications
maximum number of terminals in an analysis = 17 (mean = 2.24, median = 1)
maximum number of subtaxa (genus) = 4 (mean = 1.44, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 64
maximum number of included terminals = 17, (mean = 4.59 median = 2)
maximum number of subtaxa (genus) = 4 (mean = 2.28, median = 2)
proportion of cladograms that support basic monophyly = 0.89

Rigorous Monophyly

number of cladograms that test rigorous monophyly = 26
(Rigorous monophyly refers to cladograms with terminal species representing more than one subtaxon (genus) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 17 (mean = 7.58, median = 6)
maximum number of subtaxa (genus) = 4 (mean = 3.15, median = 3)
proportion of cladograms that support rigorous monophyly = 0.85

Pairwise Cladogram Clustering

The graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.


PCoA Ordination


NJ Tree

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

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

iTtTbiriSoTroTroSiD
171715410520
  +-------------- Megalonaias
--+                          
  |  +----------- Uniomerus  
  +--+                       
     |  +-------- Theliderma ** 
     +--+                    
        |  +----- Cyclonaias ** 
        +--+                 
           |  +-- Quadrula **   
           +--+              
              +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
1717154122218
  +-------------------------- Uniomerus           
  |                                               
--+  +----------------------- Theliderma **          
  |  |                                            
  +--+  +-------------------- Quadrula **            
     |  |                                         
     +--+  +----------------- Pustulosa_kieneriana **
        |  |                                      
        |  |              +-- Pustulosa_infucata **  
        +--+  +-----------+                       
           |  |           +-- Pustulosa_kleiniana ** 
           |  |                                   
           +--+           +-- Pustulosa_pustulosa ** 
              |  +--------+                       
              |  |        +-- Pustulosa_succissa **  
              +--+                                
                 |  +-------- Cyclonaias **          
                 +--+                             
                    |  +----- Pustulosa_necki **     
                    +--+                          
                       |  +-- Pustulosa_petrina **   
                       +--+                       
                          +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
1414124144219
                          +-- Megalonaias         
  +-----------------------+                       
  |                       +-- Uniomerus           
  |                                               
--+  +----------------------- Quadrula_quadrula **   
  |  |                                            
  |  |                    +-- Quadrula_verrucosa **  
  +--+  +-----------------+                       
     |  |                 +-- Quadrula_nobilis **    
     |  |                                         
     +--+  +----------------- Theliderma **          
        |  |                                      
        |  |           +----- Pustulosa_kieneriana **
        +--+  +--------+                          
           |  |        |  +-- Pustulosa_infucata **  
           |  |        +--+                       
           +--+           +-- Pustulosa_kleiniana ** 
              |                                   
              |  +----------- Cyclonaias **          
              |  |                                
              +--+        +-- Pustulosa_pustulosa ** 
                 |  +-----+                       
                 |  |     +-- Pustulosa_succissa **  
                 +--+                             
                    |  +----- Pustulosa_nodulata **  
                    +--+                          
                       |  +-- Pustulosa_necki **     
                       +--+                       
                          +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
1414124122219
  +----------------------------- Megalonaias         
  |                                                  
--+  +-------------------------- Uniomerus           
  |  |                                               
  +--+  +----------------------- Theliderma **          
     |  |                                            
     +--+  +-------------------- Quadrula_quadrula **   
        |  |                                         
        |  |                 +-- Quadrula_verrucosa **  
        +--+  +--------------+                       
           |  |              +-- Quadrula_nobilis **    
           |  |                                      
           |  |           +----- Pustulosa_kieneriana **
           +--+  +--------+                          
              |  |        |  +-- Pustulosa_infucata **  
              |  |        +--+                       
              |  |           +-- Pustulosa_kleiniana ** 
              +--+                                   
                 |           +-- Pustulosa_pustulosa ** 
                 |  +--------+                       
                 |  |        +-- Pustulosa_succissa **  
                 +--+                                
                    |  +-------- Cyclonaias **          
                    +--+                             
                       |  +----- Pustulosa_necki **     
                       +--+                          
                          |  +-- Pustulosa_petrina **   
                          +--+                       
                             +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
1313103161113
  +-------- Megalonaias
--+                    
  |  +----- Theliderma ** 
  +--+                 
     |  +-- Quadrula **   
     +--+              
        +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
131412311116
  +----- Megalonaias
  |                 
--+----- Theliderma ** 
  |                 
  |  +-- Quadrula **   
  +--+              
     +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
1313113161118
  +----------- Megalonaias       
  |                              
--+        +-- Quadrula_verrucosa **
  |  +-----+                     
  |  |     +-- Quadrula_nobilis **  
  +--+                           
     |  +----- Quadrula_quadrula ** 
     +--+                        
        |  +-- Theliderma **        
        +--+                     
           +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
121210442217
  +-------------------------- Megalonaias         
  |                                               
--+  +----------------------- Uniomerus           
  |  |                                            
  +--+  +-------------------- Theliderma **          
     |  |                                         
     +--+  +----------------- Quadrula **            
        |  |                                      
        |  |           +----- Pustulosa_kieneriana **
        +--+  +--------+                          
           |  |        |  +-- Pustulosa_infucata **  
           |  |        +--+                       
           |  |           +-- Pustulosa_kleiniana ** 
           +--+                                   
              |           +-- Pustulosa_pustulosa ** 
              |  +--------+                       
              |  |        +-- Pustulosa_succissa **  
              +--+                                
                 |  +-------- Cyclonaias **          
                 +--+                             
                    |  +----- Pustulosa_necki **     
                    +--+                          
                       |  +-- Pustulosa_nodulata **  
                       +--+                       
                          +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
9964932215
              +-- Megalonaias         
  +-----------+                       
  |           +-- Uniomerus           
--+                                   
  |  +----------- Theliderma **          
  |  |                                
  +--+  +-------- Quadrula **            
     |  |                             
     +--+  +----- Pustulosa_kieneriana **
        |  |                          
        |  |  +-- Cyclonaias **          
        +--+--+                       
           |  +-- Pustulosa_succissa **  
           |                          
           |  +-- Pustulosa_infucata **  
           +--+                       
              +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
9964932217
  +-------------------- Megalonaias         
  |                                         
--+  +----------------- Uniomerus           
  |  |                                      
  +--+  +-------------- Theliderma **          
     |  |                                   
     +--+  +----------- Quadrula_quadrula **   
        |  |                                
        |  |        +-- Quadrula_nobilis **    
        +--+  +-----+                       
           |  |     +-- Quadrula_verrucosa **  
           |  |                             
           +--+  +----- Pustulosa_kieneriana **
              |  |                          
              |  |  +-- Cyclonaias **          
              +--+--+                       
                 |  +-- Pustulosa_succissa **  
                 |                          
                 |  +-- Pustulosa_infucata **  
                 +--+                       
                    +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
9964642119
  +----------------- Uniomerus          
  |                                     
  |              +-- Quadrula_verrucosa ** 
--+  +-----------+                      
  |  |           +-- Quadrula_nobilis **   
  |  |                                  
  +--+  +----------- Quadrula_quadrula **  
     |  |                               
     |  +----------- Theliderma **         
     +--+                               
        |        +-- Pustulosa_infucata ** 
        |  +-----+                      
        |  |     +-- Pustulosa_kleiniana **
        +--+                            
           |  +----- Cyclonaias **         
           +--+                         
              |  +-- Pustulosa_pustulosa **
              +--+                      
                 +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
9974642118
  +-------------------- Uniomerus          
  |                                        
--+  +----------------- Theliderma **         
  |  |                                     
  +--+  +-------------- Quadrula_verrucosa ** 
     |  |                                  
     |  |           +-- Quadrula_quadrula **  
     +--+  +--------+                      
        |  |        +-- Quadrula_nobilis **   
        |  |                               
        +--+        +-- Pustulosa_infucata ** 
           |  +-----+                      
           |  |     +-- Pustulosa_kleiniana **
           +--+                            
              |  +----- Cyclonaias **         
              +--+                         
                 |  +-- Pustulosa_pustulosa **
                 +--+                      
                    +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
7753110018
  +-------------- Theliderma **          
--+                                   
  |  +----------- Quadrula_quadrula **   
  +--+                                
     |  +-------- Pustulosa_infucata **  
     +--+                             
        |  +----- Quadrula_verrucosa **  
        +--+                          
           |  +-- Pustulosa_kieneriana **
           +--+                       
              +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
6643360017
  +----------- Quadrula **           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias **         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
6644291119
  +----------------- Megalonaias        
--+                                     
  |  +-------------- Quadrula_quadrula **  
  +--+                                  
     |  +----------- Quadrula_verrucosa ** 
     +--+                               
        |  +-------- Theliderma **         
        +--+                            
           |  +----- Pustulosa_pustulosa **
           +--+                         
              |  +-- Cyclonaias **         
              +--+                      
                 +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
6643360017
  +----------- Quadrula **           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias **         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
6644321115
  +----------- Theliderma **         
--+                               
  |  +-------- Quadrula **           
  +--+                            
     |  +----- Pustulosa_pustulosa **
     +--+                         
        |  +-- Cyclonaias **         
        +--+                      
           +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
6643371117
  +-------------- Uniomerus          
  |                                  
--+           +-- Pustulosa_infucata ** 
  |  +--------+                      
  |  |        +-- Pustulosa_kleiniana **
  +--+                               
     |  +-------- Quadrula **           
     +--+                            
        |  +----- Cyclonaias **         
        +--+                         
           |  +-- Pustulosa_succissa ** 
           +--+                      
              +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
6643360017
  +----------- Quadrula **           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias **         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
5504992116
  +----- Uniomerus             
  |                            
  |  +-- Cyclonaias **            
--+  |                         
  |  +-- Pustulosa_pustulosa__1 **
  |  |                         
  |  +-- Theliderma **            
  +--+                         
     +-- Pustulosa_pustulosa__2 **
     |                         
     +-- Quadrula_quadrula **     
     |                         
     +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
5533331118
  +-------------- Uniomerus          
--+                                  
  |  +----------- Quadrula_quadrula **  
  +--+                               
     |  +-------- Quadrula_verrucosa ** 
     +--+                            
        |  +----- Pustulosa_pustulosa **
        +--+                         
           |  +-- Pustulosa_petrina **  
           +--+                      
              +-- Cyclonaias **

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

iTtTbiriSoTroTroSiD
3312110014
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
3313341116
  +----- Quadrula **  
--+                
  |  +-- Cyclonaias **
  +--+             
     +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
3312282216
  +----------- Megalonaias       
--+                              
  |  +-------- Uniomerus         
  +--+                           
     |  +----- Quadrula_verrucosa **
     +--+                        
        |  +-- Cyclonaias **        
        +--+                     
           +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
3312110014
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
3302110015
  +-- Quadrula **          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
3302110015
  +-- Quadrula **          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
29011230022
  +-- Amblema           
  |                     
  +-- Ortmanniana       
  |                     
  +-- Lampsilis         
  |                     
  +-- Cambarunio        
  |                     
--+-- Elliptio          
  |                     
  +-- Eurynia           
  |                     
  +-- Pleurobema        
  |                     
  +-- Quadrula_quadrula ** 
  |                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
22021000--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
22024000--
  +-- Pustulosa **
--+            
  +-- Quadrula **

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

iTtTbiriSoTroTroSiD
21702202230
  +-- Gonidea            
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Pleurobema         
  |                      
  +-- Unio

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

iTtTbiriSoTroTroSiD
22021000--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
22023611--
  +----- Megalonaias
--+                 
  |  +-- Cyclonaias ** 
  +--+              
     +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
22014400--
1

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

iTtTbiriSoTroTroSiD
22023122--
  +-------- Megalonaias         
--+                             
  |  +----- Quadrula_quadrula__2 **
  +--+                          
     |  +-- Quadrula_quadrula__1 **
     +--+                       
        +-- Theliderma **

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

iTtTbiriSoTroTroSiD
2202900--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
2202200--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
220119722--
1

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

iTtTbiriSoTroTroSiD
22014500--
1

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

iTtTbiriSoTroTroSiD
22022522--
  +----- Megalonaias
--+                 
  |  +-- Quadrula **   
  +--+              
     +-- Theliderma **

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

iTtTbiriSoTroTroSiD
220113700--
1

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

iTtTbiriSoTroTroSiD
21501130028
  +-------------------- Quadrula_quadrula **    
  |                                          
  +-------------------- Quadrula_verrucosa **   
  |                                          
--+                 +-- Eurynia              
  |  +--------------+                        
  |  |              +-- Pleurobema           
  |  |                                       
  +--+  +-------------- Amblema              
     |  |                                    
     |  |           +-- Obovaria             
     |  |  +--------+                        
     +--+  |        +-- Ortmanniana          
        |  |                                 
        |  |        +-- Obliquaria           
        |  +--------+                        
        +--+        +-- Lampsilis_ornata     
           |                                 
           |     +----- Lampsilis_radiata__2 
           |  +--+                           
           |  |  |  +-- Lampsilis_siliquoidea
           |  |  +--+                        
           +--+     +-- Lampsilis_radiata__1 
              |                              
              |  +----- Lampsilis_virescens  
              +--+                           
                 |  +-- Lampsilis_hydiana    
                 +--+                        
                    +-- Lampsilis_straminea

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

iTtTbiriSoTroTroSiD
22012300--
1

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

iTtTbiriSoTroTroSiD
22021900--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
2202200--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
22014400--
1

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

iTtTbiriSoTroTroSiD
22022622--
  +----- Megalonaias
--+                 
  |  +-- Quadrula **   
  +--+              
     +-- Theliderma **

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

iTtTbiriSoTroTroSiD
22011400--
1

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

iTtTbiriSoTroTroSiD
22021800--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
22021000--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
22023900--
  +-- Pustulosa **
--+            
  +-- Quadrula **

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

iTtTbiriSoTroTroSiD
220113700--
1

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

iTtTbiriSoTroTroSiD
22021900--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
220119522--
  +----- Quadrula **   
--+                 
  |  +-- Megalonaias
  +--+              
     +-- Uniomerus

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

iTtTbiriSoTroTroSiD
2202900--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
220119722--
  +----- Quadrula **   
--+                 
  |  +-- Megalonaias
  +--+              
     +-- Uniomerus

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

iTtTbiriSoTroTroSiD
2110290024
  +-------------- Quadrula **   
  |                          
--+-------------- Pustulosa **  
  |                          
  |  +----------- Pleurobema 
  +--+                       
     |  +-------- Amblema    
     +--+                    
        |  +----- Obovaria   
        +--+                 
           |  +-- Ortmanniana
           +--+              
              +-- Lampsilis

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

iTtTbiriSoTroTroSiD
22017400--
1

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

iTtTbiriSoTroTroSiD
22017800--
1

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

iTtTbiriSoTroTroSiD
2202200--
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iTtTbiriSoTroTroSiD
21002172223
  +-- Pyganodon_cataracta
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Lampsilis_straminea
--+                      
  +-- Megalonaias        
  |                      
  +-- Obliquaria         
  |                      
  +-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
22017400--
1

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

iTtTbiriSoTroTroSiD
21702202230
  +-- Gonidea            
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Aculamprotula      
  |                      
  +-- Unio

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

iTtTbiriSoTroTroSiD
22014500--
1

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013211--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11013900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015311--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11014400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017211--
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iTtTbiriSoTroTroSiD
11011800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11016700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011400--
-- Theliderma_metanevra **

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

iTtTbiriSoTroTroSiD
11013300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015611--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019011--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11018711--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11013900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015111--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11012200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017211--
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iTtTbiriSoTroTroSiD
1101900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019411--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11011100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11018411--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

Krebs, 2004 [Publ 2692], fig. 2

iTtTbiriSoTroTroSiD
1101400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011811--
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015611--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
110112000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11018200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019011--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011300--
-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
11011411--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11014500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017211--
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iTtTbiriSoTroTroSiD
11019411--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015511--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101332--
  +----- Quadrula **   
--+                 
  |  +-- Uniomerus  
  +--+              
     +-- Megalonaias

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iTtTbiriSoTroTroSiD
11011100--
-- Quadrula_quadrula **
 
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