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Page last updated
Wed 06 Nov 2024

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!

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!

Search the database pages:

Bivalvia | Unionoida | Unionoidea | Unionidae | Ambleminae | Quadrulini

subtribe QUADRULINA Ihering, 1901: cladomics
     4 genera

Cladomics Summary

number of species = 26
number of subtaxa (rank: genus) = 4

analyzed in 174 cladograms in 105 publications
maximum number of terminals in an analysis = 17, mean = 2.3
maximum number of subtaxa (genus) in an analysis = 4, mean = 1.45

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 61
maximum number of included terminals = 17, mean = 4.72
maximum number of subtaxa (genus) = 4, mean = 2.3
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 (subtribe).
maximum number of included terminals = 17, mean = 7.58
maximum number of subtaxa (genus) = 4, mean = 3.15
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

iT tT bi riS oT roT roS iD
17 17 15 4 10 5 2 0.0

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

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

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

iT tT bi riS oT roT roS iD
17 17 15 4 12 2 2 18.0 
  +-------------------------- 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

iT tT bi riS oT roT roS iD
14 14 12 4 14 4 2 19.0 
                          +-- 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

iT tT bi riS oT roT roS iD
14 14 12 4 12 2 2 19.0 
  +----------------------------- 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 **

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

iT tT bi riS oT roT roS iD
13 14 12 3 1 1 1 16.0 
  +----- Megalonaias
  |                 
--+----- Theliderma ** 
  |                 
  |  +-- Quadrula **   
  +--+              
     +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
13 13 11 3 16 1 1 18.0 
  +----------- Megalonaias       
  |                              
--+        +-- Quadrula_verrucosa **
  |  +-----+                     
  |  |     +-- Quadrula_nobilis **  
  +--+                           
     |  +----- Quadrula_quadrula ** 
     +--+                        
        |  +-- Theliderma **        
        +--+                     
           +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
13 13 10 3 16 1 1 13.0 
  +-------- Megalonaias
--+                    
  |  +----- Theliderma ** 
  +--+                 
     |  +-- Quadrula **   
     +--+              
        +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
12 12 10 4 4 2 2 17.0 
  +-------------------------- 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. 2

iT tT bi riS oT roT roS iD
9 9 6 4 93 2 2 17.0 
  +-------------------- 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

iT tT bi riS oT roT roS iD
9 9 6 4 64 2 1 19.0 
  +----------------- 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

iT tT bi riS oT roT roS iD
9 9 7 4 64 2 1 18.0 
  +-------------------- Uniomerus          
  |                                        
--+  +----------------- Theliderma **         
  |  |                                     
  +--+  +-------------- Quadrula_verrucosa ** 
     |  |                                  
     |  |           +-- Quadrula_quadrula **  
     +--+  +--------+                      
        |  |        +-- Quadrula_nobilis **   
        |  |                               
        +--+        +-- Pustulosa_infucata ** 
           |  +-----+                      
           |  |     +-- Pustulosa_kleiniana **
           +--+                            
              |  +----- Cyclonaias **         
              +--+                         
                 |  +-- Pustulosa_pustulosa **
                 +--+                      
                    +-- Pustulosa_succissa **

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

iT tT bi riS oT roT roS iD
9 9 6 4 93 2 2 15.0 
              +-- Megalonaias         
  +-----------+                       
  |           +-- Uniomerus           
--+                                   
  |  +----------- Theliderma **          
  |  |                                
  +--+  +-------- Quadrula **            
     |  |                             
     +--+  +----- Pustulosa_kieneriana **
        |  |                          
        |  |  +-- Cyclonaias **          
        +--+--+                       
           |  +-- Pustulosa_succissa **  
           |                          
           |  +-- Pustulosa_infucata **  
           +--+                       
              +-- Pustulosa_kleiniana **

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

iT tT bi riS oT roT roS iD
7 7 5 3 14 0 0 18.0 
  +-------------- Theliderma **          
--+                                   
  |  +----------- Quadrula_quadrula **   
  +--+                                
     |  +-------- Pustulosa_infucata **  
     +--+                             
        |  +----- Quadrula_verrucosa **  
        +--+                          
           |  +-- Pustulosa_kieneriana **
           +--+                       
              +-- Pustulosa_succissa **

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

iT tT bi riS oT roT roS iD
6 6 4 4 32 1 1 15.0 
  +----------- Theliderma **         
--+                               
  |  +-------- Quadrula **           
  +--+                            
     |  +----- Pustulosa_pustulosa **
     +--+                         
        |  +-- Cyclonaias **         
        +--+                      
           +-- Pustulosa_nodulata **

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

iT tT bi riS oT roT roS iD
6 6 4 3 37 1 1 17.0 
  +-------------- Uniomerus          
  |                                  
--+           +-- Pustulosa_infucata ** 
  |  +--------+                      
  |  |        +-- Pustulosa_kleiniana **
  +--+                               
     |  +-------- Quadrula **           
     +--+                            
        |  +----- Cyclonaias **         
        +--+                         
           |  +-- Pustulosa_succissa ** 
           +--+                      
              +-- Pustulosa_pustulosa **

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

iT tT bi riS oT roT roS iD
6 6 4 3 36 0 0 17.0 
  +----------- Quadrula **           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias **         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iT tT bi riS oT roT roS iD
6 6 4 3 36 0 0 17.0 
  +----------- Quadrula **           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias **         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iT tT bi riS oT roT roS iD
6 6 4 3 36 0 0 17.0 
  +----------- Quadrula **           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias **         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iT tT bi riS oT roT roS iD
6 6 4 4 29 1 1 19.0 
  +----------------- Megalonaias        
--+                                     
  |  +-------------- Quadrula_quadrula **  
  +--+                                  
     |  +----------- Quadrula_verrucosa ** 
     +--+                               
        |  +-------- Theliderma **         
        +--+                            
           |  +----- Pustulosa_pustulosa **
           +--+                         
              |  +-- Cyclonaias **         
              +--+                      
                 +-- Pustulosa_nodulata **

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

iT tT bi riS oT roT roS iD
5 5 0 4 99 2 1 16.0 
  +----- Uniomerus             
  |                            
  |  +-- Cyclonaias **            
--+  |                         
  |  +-- Pustulosa_pustulosa__1 **
  |  |                         
  |  +-- Theliderma **            
  +--+                         
     +-- Pustulosa_pustulosa__2 **
     |                         
     +-- Quadrula_quadrula **     
     |                         
     +-- Quadrula_verrucosa **

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

iT tT bi riS oT roT roS iD
5 5 3 3 33 1 1 18.0 
  +-------------- Uniomerus          
--+                                  
  |  +----------- Quadrula_quadrula **  
  +--+                               
     |  +-------- Quadrula_verrucosa ** 
     +--+                            
        |  +----- Pustulosa_pustulosa **
        +--+                         
           |  +-- Pustulosa_petrina **  
           +--+                      
              +-- Cyclonaias **

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

iT tT bi riS oT roT roS iD
3 3 0 2 11 0 0 15.0 
  +-- Quadrula **          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

iT tT bi riS oT roT roS iD
3 3 1 3 34 1 1 16.0 
  +----- Quadrula **  
--+                
  |  +-- Cyclonaias **
  +--+             
     +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
3 3 1 2 11 0 0 14.0 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
3 3 1 2 11 0 0 14.0 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
3 3 0 2 11 0 0 15.0 
  +-- Quadrula **          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

iT tT bi riS oT roT roS iD
3 3 1 2 32 2 2 16.0 
  +----------- Megalonaias       
--+                              
  |  +-------- Uniomerus         
  +--+                           
     |  +----- Quadrula_verrucosa **
     +--+                        
        |  +-- Cyclonaias **        
        +--+                     
           +-- Quadrula_quadrula **

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

iT tT bi riS oT roT roS iD
2 2 0 2 40 0 0 -- 
  +-- Pustulosa **
--+            
  +-- Quadrula **

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

iT tT bi riS oT roT roS iD
2 17 0 2 20 2 2 30.0 
  +-- Gonidea            
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Pleurobema         
  |                      
  +-- Unio

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

iT tT bi riS oT roT roS iD
2 2 0 1 196 2 2 -- 
  +----- Quadrula **   
--+                 
  |  +-- Megalonaias
  +--+              
     +-- Uniomerus

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

iT tT bi riS oT roT roS iD
2 2 0 1 194 2 2 -- 
  +----- Quadrula **   
--+                 
  |  +-- Megalonaias
  +--+              
     +-- Uniomerus

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

iT tT bi riS oT roT roS iD
2 2 0 1 78 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 2 18 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 2 19 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 2 9 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 2 2 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 2 31 2 2 -- 
  +-------- Megalonaias         
--+                             
  |  +----- Quadrula_quadrula__2 **
  +--+                          
     |  +-- Quadrula_quadrula__1 **
     +--+                       
        +-- Theliderma **

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

iT tT bi riS oT roT roS iD
2 2 0 1 44 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 1 23 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 15 0 1 13 0 0 28.0 
  +-------------------- 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

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

iT tT bi riS oT roT roS iD
2 2 0 2 26 2 2 -- 
  +----- Megalonaias
--+                 
  |  +-- Quadrula **   
  +--+              
     +-- Theliderma **

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

iT tT bi riS oT roT roS iD
2 2 0 1 196 2 2 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 1 45 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 2 39 0 0 -- 
  +-- Pustulosa **
--+            
  +-- Quadrula **

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

iT tT bi riS oT roT roS iD
2 17 0 2 20 2 2 30.0 
  +-- Gonidea            
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Pyganodon_cataracta
  |                      
  +-- Lampsilis_straminea
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Megalonaias        
  |                      
  +-- Obliquaria         
  |                      
--+-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus          
  |                      
  +-- Amblema            
  |                      
  +-- Elliptio           
  |                      
  +-- Fusconaia          
  |                      
  +-- Plectomerus        
  |                      
  +-- Aculamprotula      
  |                      
  +-- Unio

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

iT tT bi riS oT roT roS iD
2 2 0 1 14 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 10 0 2 17 2 2 23.0 
  +-- Pyganodon_cataracta
  |                      
  +-- Pyganodon_grandis  
  |                      
  +-- Utterbackia        
  |                      
  +-- Lampsilis_teres    
  |                      
  +-- Lampsilis_straminea
--+                      
  +-- Megalonaias        
  |                      
  +-- Obliquaria         
  |                      
  +-- Theliderma **         
  |                      
  +-- Quadrula **           
  |                      
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
2 11 0 2 9 0 0 24.0 
  +-------------- Quadrula **   
  |                          
--+-------------- Pustulosa **  
  |                          
  |  +----------- Pleurobema 
  +--+                       
     |  +-------- Amblema    
     +--+                    
        |  +----- Obovaria   
        +--+                 
           |  +-- Ortmanniana
           +--+              
              +-- Lampsilis

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

iT tT bi riS oT roT roS iD
2 2 0 2 19 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 1 137 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 1 74 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 2 2 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 2 10 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 1 137 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 9 0 1 123 0 0 22.0 
  +-- Amblema           
  |                     
  +-- Ortmanniana       
  |                     
  +-- Lampsilis         
  |                     
  +-- Cambarunio        
  |                     
--+-- Elliptio          
  |                     
  +-- Eurynia           
  |                     
  +-- Pleurobema        
  |                     
  +-- Quadrula_quadrula ** 
  |                     
  +-- Quadrula_verrucosa **

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

iT tT bi riS oT roT roS iD
2 2 0 1 45 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 2 2 0 0 -- 
  +-- Quadrula ** 
--+            
  +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
2 2 0 2 25 2 2 -- 
  +----- Megalonaias
--+                 
  |  +-- Quadrula **   
  +--+              
     +-- Theliderma **

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

iT tT bi riS oT roT roS iD
2 2 0 1 44 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 1 74 0 0 -- 
-- (Quadrula

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

iT tT bi riS oT roT roS iD
2 2 0 2 36 1 1 -- 
  +----- Megalonaias
--+                 
  |  +-- Cyclonaias ** 
  +--+              
     +-- Pustulosa **

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

iT tT bi riS oT roT roS iD
1 1 0 1 82 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 90 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 22 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 95 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 14 0 0 -- 
-- Theliderma_metanevra

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

iT tT bi riS oT roT roS iD
1 1 0 1 11 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 68 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 9 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 31 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 33 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 1 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 18 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 5 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 42 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 17 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 32 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 95 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 14 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 10 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 56 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 13 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 6 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 48 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 51 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 32 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 27 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 11 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 3 3 2 -- 
  +----- Quadrula **   
--+                 
  |  +-- Uniomerus  
  +--+              
     +-- Megalonaias

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

iT tT bi riS oT roT roS iD
1 1 0 1 39 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 5 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 38 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 21 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 33 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 91 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 15 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 24 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 6 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 6 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 16 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 31 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 119 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 91 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 19 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 73 1 1 -- 
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iT tT bi riS oT roT roS iD
1 1 0 1 39 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 15 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 12 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 35 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 53 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 19 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 21 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 5 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 73 1 1 -- 
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iT tT bi riS oT roT roS iD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 56 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 24 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 29 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 31 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 10 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 49 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 13 0 0 -- 
-- Quadrula_nobilis

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

iT tT bi riS oT roT roS iD
1 1 0 1 44 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 12 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 21 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 38 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 27 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 5 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 22 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 90 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 11 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 52 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 9 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 39 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 19 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 39 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 8 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 46 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 73 1 1 -- 
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iT tT bi riS oT roT roS iD
1 1 0 1 33 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 18 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 14 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 52 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 16 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 24 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 35 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 35 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 6 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 19 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 35 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 6 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 8 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 18 1 1 -- 
  +-- Uniomerus
--+            
  +-- Quadrula **

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

iT tT bi riS oT roT roS iD
1 1 0 1 56 1 1 -- 
  +-- Quadrula ** 
--+            
  +-- Uniomerus

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

iT tT bi riS oT roT roS iD
1 1 0 1 20 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 28 0 0 -- 
-- Quadrula_quadrula

Krebs, 2004 [Publ 2692], fig. 2

iT tT bi riS oT roT roS iD
1 1 0 1 4 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 44 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 16 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 18 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 29 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula

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

iT tT bi riS oT roT roS iD
1 1 0 1 25 0 0 -- 
-- Quadrula_quadrula

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