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Page last updated
Thu 3 September 2025

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

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

Search the database pages:

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

genus Quadrula Rafinesque, 1820: cladomics
     5 Recent valid species (9 total species)

Cladomics Summary

number of species = 5 (9 total)
analyzed in 191 cladograms in 115 publications
maximum number of terminals in an analysis = 4 (mean = 1.26, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 35
maximum number of included terminals = 4, (mean = 2.4 median = 2)
proportion of cladograms that support basic monophyly = 0.6

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

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

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
45346281230
                 +-- Theliderma_johnsoni  
        +--------+                        
        |        +-- Theliderma_metanevra 
        |                                 
  +-----+        +-- Quadrula_nobilis **     
  |     |  +-----+                        
  |     |  |     +-- Quadrula_verrucosa **   
  |     +--+                              
  |        |  +----- Theliderma_cylindrica
  |        +--+                           
  |           |  +-- Quadrula_fragosa **     
--+           +--+                        
  |              +-- Quadrula_quadrula **    
  |                                       
  |           +----- 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. 3

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
3313261432
  +----------------------- Theliderma          
  |                                            
  |                 +----- Quadrula_quadrula **   
--+  +--------------+                          
  |  |              |  +-- Quadrula_nobilis **    
  |  |              +--+                       
  |  |                 +-- Quadrula_verrucosa **  
  +--+                                         
     |  +----------------- Pustulosa_kieneriana
     |  |                                      
     |  |              +-- Pustulosa_infucata  
     +--+  +-----------+                       
        |  |           +-- Pustulosa_kleiniana 
        |  |                                   
        +--+           +-- Pustulosa_pustulosa 
           |  +--------+                       
           |  |        +-- Pustulosa_succissa  
           +--+                                
              |  +-------- Cyclonaias          
              +--+                             
                 |  +----- Pustulosa_necki     
                 +--+                          
                    |  +-- Pustulosa_petrina   
                    +--+                       
                       +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
4424231333
  +----------------- Theliderma        
--+                                    
  |  +-------------- Cyclonaias        
  +--+                                 
     |  +----------- Pustulosa         
     +--+                              
        |  +-------- Quadrula_nobilis **  
        +--+                           
           |  +----- Quadrula_verrucosa **
           +--+                        
              |  +-- Quadrula_fragosa **  
              +--+                     
                 +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
385399637
  +-------------- Theliderma          
  |                                   
--+  +----------- Quadrula_quadrula **   
  |  |                                
  |  |        +-- Quadrula_nobilis **    
  +--+  +-----+                       
     |  |     +-- Quadrula_verrucosa **  
     |  |                             
     +--+  +----- Pustulosa_kieneriana
        |  |                          
        |  |  +-- Cyclonaias          
        +--+--+                       
           |  +-- Pustulosa_succissa  
           |                          
           |  +-- Pustulosa_infucata  
           +--+                       
              +-- Pustulosa_kleiniana

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

iTtTbiriSoTroTroSiD
396370637
              +-- Quadrula_verrucosa ** 
  +-----------+                      
  |           +-- Quadrula_nobilis **   
  |                                  
--+  +----------- Quadrula_quadrula **  
  |  |                               
  |  +----------- Theliderma         
  +--+                               
     |        +-- Pustulosa_infucata 
     |  +-----+                      
     |  |     +-- Pustulosa_kleiniana
     +--+                            
        |  +----- Cyclonaias         
        +--+                         
           |  +-- Pustulosa_pustulosa
           +--+                      
              +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
3324111025
  +----------- Pustulosa           
--+                                
  |  +-------- Quadrula_quadrula__1 **
  +--+                             
     |  +----- Quadrula_verrucosa **  
     +--+                          
        |  +-- Quadrula_nobilis **    
        +--+                       
           +-- Quadrula_quadrula__2 **

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

iTtTbiriSoTroTroSiD
3313261024
  +----------- Theliderma        
--+                              
  |  +-------- Pustulosa         
  +--+                           
     |  +----- Quadrula_verrucosa **
     +--+                        
        |  +-- Quadrula_quadrula ** 
        +--+                     
           +-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
3141232511314
  +----------------------- Quadrula_quadrula **   
  |                                            
  |                    +-- Quadrula_verrucosa **  
--+  +-----------------+                       
  |  |                 +-- Quadrula_nobilis **    
  |  |                                         
  +--+  +----------------- Theliderma          
     |  |                                      
     |  |           +----- Pustulosa_kieneriana
     +--+  +--------+                          
        |  |        |  +-- Pustulosa_infucata  
        |  |        +--+                       
        +--+           +-- Pustulosa_kleiniana 
           |                                   
           |  +----------- Cyclonaias          
           |  |                                
           +--+        +-- Pustulosa_pustulosa 
              |  +-----+                       
              |  |     +-- Pustulosa_succissa  
              +--+                             
                 |  +----- Pustulosa_nodulata  
                 +--+                          
                    |  +-- Pustulosa_necki     
                    +--+                       
                       +-- Pustulosa_petrina

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

iTtTbiriSoTroTroSiD
3121032311311
  +----------------------- Theliderma          
  |                                            
--+  +-------------------- Quadrula_quadrula **   
  |  |                                         
  |  |                 +-- Quadrula_verrucosa **  
  +--+  +--------------+                       
     |  |              +-- Quadrula_nobilis **    
     |  |                                      
     |  |           +----- Pustulosa_kieneriana
     +--+  +--------+                          
        |  |        |  +-- Pustulosa_infucata  
        |  |        +--+                       
        |  |           +-- Pustulosa_kleiniana 
        +--+                                   
           |           +-- Pustulosa_pustulosa 
           |  +--------+                       
           |  |        +-- Pustulosa_succissa  
           +--+                                
              |  +-------- Cyclonaias          
              +--+                             
                 |  +----- Pustulosa_necki     
                 +--+                          
                    |  +-- Pustulosa_petrina   
                    +--+                       
                       +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
3131132610210
        +-- Quadrula_verrucosa **
  +-----+                     
  |     +-- Quadrula_nobilis **  
--+                           
  |  +----- Quadrula_quadrula ** 
  +--+                        
     |  +-- Theliderma        
     +--+                     
        +-- Pustulosa

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

iTtTbiriSoTroTroSiD
386370639
  +----------------- 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
331399634
  +----------- Theliderma          
  |                                
  |     +----- Quadrula_verrucosa **  
--+  +--+                          
  |  |  |  +-- Quadrula_quadrula **   
  |  |  +--+                       
  |  |     +-- Quadrula_nobilis **    
  +--+                             
     |  +----- Pustulosa_kieneriana
     |  |                          
     |  |  +-- Cyclonaias          
     +--+--+                       
        |  +-- Pustulosa_succissa  
        |                          
        |  +-- Pustulosa_infucata  
        +--+                       
           +-- Pustulosa_kleiniana

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

iTtTbiriSoTroTroSiD
22024400--
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
2502103337
  +-- Cyclonaias            
  |                         
  +-- Pustulosa_pustulosa__1
  |                         
  +-- Theliderma            
--+                         
  +-- Pustulosa_pustulosa__2
  |                         
  +-- Quadrula_quadrula **     
  |                         
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
220219800--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
22027800--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
220219600--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
22024500--
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
22024400--
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
220219800--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
260233438
  +-------------- Quadrula_quadrula **  
--+                                  
  |  +----------- Quadrula_verrucosa ** 
  +--+                               
     |  +-------- Theliderma         
     +--+                            
        |  +----- Pustulosa_pustulosa
        +--+                         
           |  +-- Cyclonaias         
           +--+                      
              +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
220213600--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
22024500--
  +-- Quadrula_verrucosa **
--+                     
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
22023643--
  +----------- Theliderma         
  |                               
--+        +-- Quadrula_verrucosa ** 
  |  +-----+                      
  |  |     +-- Quadrula_quadrula **  
  +--+                            
     |  +----- Pustulosa_pustulosa
     +--+                         
        |  +-- Cyclonaias         
        +--+                      
           +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
220214103--
  +-------------------- Theliderma          
  |                                         
  |                 +-- Quadrula_quadrula **   
--+  +--------------+                       
  |  |              +-- Quadrula_verrucosa **  
  |  |                                      
  |  |           +----- Pustulosa_kieneriana
  +--+  +--------+                          
     |  |        |  +-- Pustulosa_infucata  
     |  |        +--+                       
     |  |           +-- Pustulosa_kleiniana 
     +--+                                   
        |           +-- Pustulosa_pustulosa 
        |  +--------+                       
        |  |        +-- Pustulosa_succissa  
        +--+                                
           |  +-------- Cyclonaias          
           +--+                             
              |  +----- Pustulosa_necki     
              +--+                          
                 |  +-- Pustulosa_nodulata  
                 +--+                       
                    +-- Pustulosa_petrina

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

iTtTbiriSoTroTroSiD
22031400--
  +----- Quadrula_quadrula__2 **
--+                          
  |  +-- Quadrula_quadrula__1 **
  +--+                       
     +-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
220213600--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
21502130017
  +-------------------- Quadrula_quadrula **    
  |                                          
  +-------------------- Quadrula_verrucosa **   
  |                                          
--+                 +-- Eurynia              
  |  +--------------+                        
  |  |              +-- Pleurobema           
  |  |                                       
  +--+  +-------------- Amblema              
     |  |                                    
     |  |           +-- Obovaria             
     |  |  +--------+                        
     +--+  |        +-- Ortmanniana          
        |  |                                 
        |  |        +-- Obliquaria           
        |  +--------+                        
        +--+        +-- Lampsilis_ornata     
           |                                 
           |     +----- Lampsilis_radiata__2 
           |  +--+                           
           |  |  |  +-- Lampsilis_siliquoidea
           |  |  +--+                        
           +--+     +-- Lampsilis_radiata__1 
              |                              
              |  +----- Lampsilis_virescens  
              +--+                           
                 |  +-- Lampsilis_hydiana    
                 +--+                        
                    +-- Lampsilis_straminea

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

iTtTbiriSoTroTroSiD
22022300--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
29021220010
  +-- Amblema           
  |                     
  +-- Ortmanniana       
  |                     
  +-- Lampsilis         
  |                     
  +-- Cambarunio        
  |                     
--+-- Elliptio          
  |                     
  +-- Eurynia           
  |                     
  +-- Pleurobema        
  |                     
  +-- Quadrula_quadrula ** 
  |                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
22027400--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
250236326
  +----------- Quadrula_quadrula **  
--+                               
  |  +-------- Quadrula_verrucosa ** 
  +--+                            
     |  +----- Pustulosa_pustulosa
     +--+                         
        |  +-- Pustulosa_petrina  
        +--+                      
           +-- Cyclonaias

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

iTtTbiriSoTroTroSiD
22027400--
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
250216526
  +-------------- Theliderma          
--+                                   
  |  +----------- Quadrula_quadrula **   
  +--+                                
     |  +-------- Pustulosa_infucata  
     +--+                             
        |  +----- Quadrula_verrucosa **  
        +--+                          
           |  +-- Pustulosa_kieneriana
           +--+                       
              +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
230229114
  +----- Quadrula_verrucosa **
--+                        
  |  +-- Cyclonaias        
  +--+                     
     +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014152--
  +----------- Quadrula_quadrula **  
  |                               
--+        +-- Pustulosa_infucata 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
11012200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11018100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012111--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011321--
  +-- Quadrula_nobilis **
--+                   
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11011011--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
1101311--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11015500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012011--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11011100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013100--
-- Quadrula_quadrula **

Klymus et al., 2025 [Publ 4814], fig. 3B

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011011--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11011600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013800--
-- Quadrula_quadrula **

Klymus et al., 2025 [Publ 4814], fig. 3A

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012611--
  +-- Quadrula_quadrula **
--+                    
  +-- Theliderma

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

iTtTbiriSoTroTroSiD
11011000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11016800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019100--
-- Quadrula_quadrula **

Campbell & Lydeard, 2026 [Publ 4810], fig. 3_5

iTtTbiriSoTroTroSiD
110114000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011321--
  +-- Quadrula_nobilis **
--+                   
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11011111--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11014111--
  +-- Pustulosa        
--+                    
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Pustulosa        
--+                    
  +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101311--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11015200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011911--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11013200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
12023211--
  +----- Quadrula_quadrula__2 **
--+                          
  |  +-- Quadrula_quadrula__1 **
  +--+                       
     +-- Theliderma

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

iTtTbiriSoTroTroSiD
110112100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012711--
  +-- Quadrula_quadrula **
--+                    
  +-- Theliderma

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011300--
-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
11014400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012111--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011111--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11011321--
  +-- Quadrula_nobilis **  
  |                     
--+-- Pustulosa_infucata
  |                     
  +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
11014400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011011--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11018700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014152--
  +----------- Quadrula_quadrula **  
  |                               
--+        +-- Pustulosa_infucata 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
1101900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014152--
  +----------- Quadrula_quadrula **  
  |                               
--+        +-- Pustulosa_infucata 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
11015200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013722--
  +----- Quadrula_quadrula **
--+                       
  |  +-- Cyclonaias       
  +--+                    
     +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11012011--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11019000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11018500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014252--
           +-- Pustulosa_infucata 
  +--------+                      
  |        +-- Pustulosa_kleiniana
--+                               
  |  +-------- Quadrula_quadrula **  
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa 
        +--+                      
           +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
11013600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11019500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101500--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011321--
  +-- Quadrula_nobilis **  
  |                     
--+-- Pustulosa_infucata
  |                     
  +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
11015600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012400--
-- Quadrula_quadrula **

Campbell & Lydeard, 2026 [Publ 4810], fig. 4

iTtTbiriSoTroTroSiD
11014600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014800--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011811--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011800--
-- Quadrula_quadrula **

Krebs, 2004 [Publ 2692], fig. 2

iTtTbiriSoTroTroSiD
1101400--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11017300--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11015200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011600--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101311--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11011111--
  +-- Quadrula_quadrula **
--+                    
  +-- Pustulosa

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

iTtTbiriSoTroTroSiD
11019000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014700--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011900--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11011100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11012200--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
1101100--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11014000--
-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
11013500--
-- Quadrula_quadrula **
 
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