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Data last updated
Mon 1 Dec 2025

Bivalvia | Unionoida | 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 184 cladograms in 112 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. 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

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 **

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. 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. 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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 2 74 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 **

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 **

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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. 3B-M

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 24 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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 7 0 0 -- 
-- 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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 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. 4

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Krebs, 2004 [Publ 2692], fig. 2

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 6 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

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

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

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

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

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

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

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 **

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 **

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 43 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., 2023 [Publ 4302], fig. 1

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 87 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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 121 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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 68 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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 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 **

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

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

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

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

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

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

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

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