Bivalvia | Unionida | Unionoidea | Unionidae | Ambleminae | Quadrulini | Quadrulina

genus Quadrula Rafinesque, 1820: cladomics

5 Recent valid species (9 total species)

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 = 5 (9 total)
analyzed in 186 cladograms in 113 publications
maximum number of terminals in an analysis = 4 (mean = 1.25, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 34
maximum number of included terminals = 4, (mean = 2.35 median = 2)
proportion of cladograms that support basic monophyly = 0.62

Pairwise Cladogram Clustering

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

PCoA Ordination of Quadrula cladogram distances
PCoA Ordination

NJ Tree of Quadrula cladogram distances
NJ Tree

Cladograms

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

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
4 4 2 4 23 13 3 0 
  +----------------- Theliderma        
--+                                    
  |  +-------------- Cyclonaias        
  +--+                                 
     |  +----------- Pustulosa         
     +--+                              
        |  +-------- Quadrula_nobilis **  
        +--+                           
           |  +----- Quadrula_verrucosa **
           +--+                        
              |  +-- Quadrula_fragosa **  
              +--+                     
                 +-- Quadrula_quadrula **

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
3 3 1 3 26 14 3 3 
  +----------------------- Theliderma          
  |                                            
  |                 +----- Quadrula_quadrula **   
--+  +--------------+                          
  |  |              |  +-- Quadrula_nobilis **    
  |  |              +--+                       
  |  |                 +-- Quadrula_verrucosa **  
  +--+                                         
     |  +----------------- Pustulosa_kieneriana
     |  |                                      
     |  |              +-- Pustulosa_infucata  
     +--+  +-----------+                       
        |  |           +-- Pustulosa_kleiniana 
        |  |                                   
        +--+           +-- Pustulosa_pustulosa 
           |  +--------+                       
           |  |        +-- Pustulosa_succissa  
           +--+                                
              |  +-------- Cyclonaias          
              +--+                             
                 |  +----- Pustulosa_necki     
                 +--+                          
                    |  +-- Pustulosa_petrina   
                    +--+                       
                       +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
3 9 6 3 70 6 3 9 
              +-- Quadrula_verrucosa ** 
  +-----------+                      
  |           +-- Quadrula_nobilis **   
  |                                  
--+  +----------- Quadrula_quadrula **  
  |  |                               
  |  +----------- Theliderma         
  +--+                               
     |        +-- Pustulosa_infucata 
     |  +-----+                      
     |  |     +-- Pustulosa_kleiniana
     +--+                            
        |  +----- Cyclonaias         
        +--+                         
           |  +-- Pustulosa_pustulosa
           +--+                      
              +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
3 12 10 3 23 11 3 12 
  +----------------------- Theliderma          
  |                                            
--+  +-------------------- Quadrula_quadrula **   
  |  |                                         
  |  |                 +-- Quadrula_verrucosa **  
  +--+  +--------------+                       
     |  |              +-- Quadrula_nobilis **    
     |  |                                      
     |  |           +----- Pustulosa_kieneriana
     +--+  +--------+                          
        |  |        |  +-- Pustulosa_infucata  
        |  |        +--+                       
        |  |           +-- Pustulosa_kleiniana 
        +--+                                   
           |           +-- Pustulosa_pustulosa 
           |  +--------+                       
           |  |        +-- Pustulosa_succissa  
           +--+                                
              |  +-------- Cyclonaias          
              +--+                             
                 |  +----- Pustulosa_necki     
                 +--+                          
                    |  +-- Pustulosa_petrina   
                    +--+                       
                       +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
3 3 1 3 99 6 3 3 
  +----------- Theliderma          
  |                                
  |     +----- Quadrula_verrucosa **  
--+  +--+                          
  |  |  |  +-- Quadrula_quadrula **   
  |  |  +--+                       
  |  |     +-- Quadrula_nobilis **    
  +--+                             
     |  +----- Pustulosa_kieneriana
     |  |                          
     |  |  +-- Cyclonaias          
     +--+--+                       
        |  +-- Pustulosa_succissa  
        |                          
        |  +-- Pustulosa_infucata  
        +--+                       
           +-- Pustulosa_kleiniana

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

iTtTbiriSoTroTroSiD
3 3 1 3 26 10 2 3 
  +----------- Theliderma        
--+                              
  |  +-------- Pustulosa         
  +--+                           
     |  +----- Quadrula_verrucosa **
     +--+                        
        |  +-- Quadrula_quadrula ** 
        +--+                     
           +-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
3 13 11 3 26 10 2 13 
        +-- Quadrula_verrucosa **
  +-----+                     
  |     +-- Quadrula_nobilis **  
--+                           
  |  +----- Quadrula_quadrula ** 
  +--+                        
     |  +-- Theliderma        
     +--+                     
        +-- Pustulosa

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

iTtTbiriSoTroTroSiD
3 14 12 3 25 11 3 14 
  +----------------------- Quadrula_quadrula **   
  |                                            
  |                    +-- Quadrula_verrucosa **  
--+  +-----------------+                       
  |  |                 +-- Quadrula_nobilis **    
  |  |                                         
  +--+  +----------------- Theliderma          
     |  |                                      
     |  |           +----- Pustulosa_kieneriana
     +--+  +--------+                          
        |  |        |  +-- Pustulosa_infucata  
        |  |        +--+                       
        +--+           +-- Pustulosa_kleiniana 
           |                                   
           |  +----------- Cyclonaias          
           |  |                                
           +--+        +-- Pustulosa_pustulosa 
              |  +-----+                       
              |  |     +-- Pustulosa_succissa  
              +--+                             
                 |  +----- Pustulosa_nodulata  
                 +--+                          
                    |  +-- Pustulosa_necki     
                    +--+                       
                       +-- Pustulosa_petrina

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

iTtTbiriSoTroTroSiD
3 8 6 3 70 6 3 8 
  +----------------- Theliderma         
  |                                     
--+  +-------------- Quadrula_verrucosa ** 
  |  |                                  
  |  |           +-- Quadrula_quadrula **  
  +--+  +--------+                      
     |  |        +-- Quadrula_nobilis **   
     |  |                               
     +--+        +-- Pustulosa_infucata 
        |  +-----+                      
        |  |     +-- Pustulosa_kleiniana
        +--+                            
           |  +----- Cyclonaias         
           +--+                         
              |  +-- Pustulosa_pustulosa
              +--+                      
                 +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
3 3 2 4 11 10 2 4 
  +----------- Pustulosa           
--+                                
  |  +-------- Quadrula_quadrula__1 **
  +--+                             
     |  +----- Quadrula_verrucosa **  
     +--+                          
        |  +-- Quadrula_nobilis **    
        +--+                       
           +-- Quadrula_quadrula__2 **

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

iTtTbiriSoTroTroSiD
3 8 5 3 99 6 3 8 
  +-------------- Theliderma          
  |                                   
--+  +----------- Quadrula_quadrula **   
  |  |                                
  |  |        +-- Quadrula_nobilis **    
  +--+  +-----+                       
     |  |     +-- Quadrula_verrucosa **  
     |  |                             
     +--+  +----- Pustulosa_kieneriana
        |  |                          
        |  |  +-- Cyclonaias          
        +--+--+                       
           |  +-- Pustulosa_succissa  
           |                          
           |  +-- Pustulosa_infucata  
           +--+                       
              +-- Pustulosa_kleiniana

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

iTtTbiriSoTroTroSiD
2 2 0 2 199 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 5 0 2 36 3 2 5 
  +----------- Quadrula_quadrula **  
--+                               
  |  +-------- Quadrula_verrucosa ** 
  +--+                            
     |  +----- Pustulosa_pustulosa
     +--+                         
        |  +-- Pustulosa_petrina  
        +--+                      
           +-- Cyclonaias

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

iTtTbiriSoTroTroSiD
2 2 0 2 45 0 0 -- 
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
2 2 0 2 197 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 6 0 2 33 4 3 6 
  +-------------- Quadrula_quadrula **  
--+                                  
  |  +----------- Quadrula_verrucosa ** 
  +--+                               
     |  +-------- Theliderma         
     +--+                            
        |  +----- Pustulosa_pustulosa
        +--+                         
           |  +-- Cyclonaias         
           +--+                      
              +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
2 3 0 2 29 1 1 3 
  +----- Quadrula_verrucosa **
--+                        
  |  +-- Cyclonaias        
  +--+                     
     +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
2 2 0 2 137 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 137 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 74 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 14 10 3 -- 
  +-------------------- Theliderma          
  |                                         
  |                 +-- Quadrula_quadrula **   
--+  +--------------+                       
  |  |              +-- Quadrula_verrucosa **  
  |  |                                      
  |  |           +----- Pustulosa_kieneriana
  +--+  +--------+                          
     |  |        |  +-- Pustulosa_infucata  
     |  |        +--+                       
     |  |           +-- Pustulosa_kleiniana 
     +--+                                   
        |           +-- Pustulosa_pustulosa 
        |  +--------+                       
        |  |        +-- Pustulosa_succissa  
        +--+                                
           |  +-------- Cyclonaias          
           +--+                             
              |  +----- Pustulosa_necki     
              +--+                          
                 |  +-- Pustulosa_nodulata  
                 +--+                       
                    +-- Pustulosa_petrina

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

iTtTbiriSoTroTroSiD
2 2 0 2 74 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 3 14 0 0 -- 
  +----- Quadrula_quadrula__2 **
--+                          
  |  +-- Quadrula_quadrula__1 **
  +--+                       
     +-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
2 15 0 2 13 0 0 16 
  +-------------------- 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

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

iTtTbiriSoTroTroSiD
2 2 0 2 45 0 0 -- 
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
2 2 0 2 44 0 0 -- 
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
2 2 0 2 199 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 5 0 2 102 3 3 6 
  +-- Cyclonaias            
  |                         
  +-- Pustulosa_pustulosa__1
  |                         
  +-- Theliderma            
--+                         
  +-- Pustulosa_pustulosa__2
  |                         
  +-- Quadrula_quadrula **     
  |                         
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 23 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 44 0 0 -- 
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
2 9 0 2 123 0 0 9 
  +-- Amblema           
  |                     
  +-- Ortmanniana       
  |                     
  +-- Lampsilis         
  |                     
  +-- Cambarunio        
  |                     
--+-- Elliptio          
  |                     
  +-- Eurynia           
  |                     
  +-- Pleurobema        
  |                     
  +-- Quadrula_quadrula ** 
  |                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 36 4 3 -- 
  +----------- Theliderma         
  |                               
--+        +-- Quadrula_verrucosa ** 
  |  +-----+                      
  |  |     +-- Quadrula_quadrula **  
  +--+                            
     |  +----- Pustulosa_pustulosa
     +--+                         
        |  +-- Cyclonaias         
        +--+                      
           +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
2 2 0 2 78 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 5 0 2 16 5 2 5 
  +-------------- Theliderma          
--+                                   
  |  +----------- Quadrula_quadrula **   
  +--+                                
     |  +-------- Pustulosa_infucata  
     +--+                             
        |  +----- Quadrula_verrucosa **  
        +--+                          
           |  +-- Pustulosa_kieneriana
           +--+                       
              +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 73 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 91 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 36 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 27 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
1 1 0 1 55 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 31 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 43 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 26 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Theliderma

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

iTtTbiriSoTroTroSiD
1 1 0 1 8 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 2 1 -- 
  +-- Quadrula_nobilis **
--+                   
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 1 1 -- 
  +-- Pustulosa        
--+                    
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 5 2 -- 
  +----------- Quadrula_quadrula **  
  |                               
--+        +-- Pustulosa_infucata 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 15 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 121 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 90 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 39 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 8 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 91 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 2 0 2 32 1 1 -- 
  +----- Quadrula_quadrula__2 **
--+                          
  |  +-- Quadrula_quadrula__1 **
  +--+                       
     +-- Theliderma

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 75 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 87 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 9 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 68 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 15 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 90 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 1 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 5 2 -- 
  +----------- Quadrula_quadrula **  
  |                               
--+        +-- Pustulosa_infucata 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 47 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 0 0 -- 
-- Quadrula_quadrula **

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 36 2 2 -- 
  +----- Quadrula_quadrula **
--+                       
  |  +-- Cyclonaias       
  +--+                    
     +-- Pustulosa

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 85 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 42 5 2 -- 
           +-- Pustulosa_infucata 
  +--------+                      
  |        +-- Pustulosa_kleiniana
--+                               
  |  +-------- Quadrula_quadrula **  
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 14 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 15 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 2 1 -- 
  +-- Quadrula_nobilis **  
  |                     
--+-- Pustulosa_infucata
  |                     
  +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 53 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 32 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 17 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 2 1 -- 
  +-- Quadrula_nobilis **
--+                   
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 37 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 48 0 0 -- 
-- Quadrula_quadrula **

Krebs, 2004 [Publ 2692], fig. 2

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 18 1 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 43 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 27 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Theliderma

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

iTtTbiriSoTroTroSiD
1 1 0 1 56 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 39 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 35 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 27 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 1 1 -- 
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 73 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 1 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 95 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Pustulosa        
--+                    
  +-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 2 1 -- 
  +-- Quadrula_nobilis **  
  |                     
--+-- Pustulosa_infucata
  |                     
  +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 5 2 -- 
  +----------- Quadrula_quadrula **  
  |                               
--+        +-- Pustulosa_infucata 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 73 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 95 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 82 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 33 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 1 1 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 56 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 38 0 0 -- 
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 0 0 -- 
-- Quadrula_quadrula **