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 189 cladograms in 114 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 of Quadrula cladogram distances
PCoA Ordination

NJ Tree of Quadrula cladogram distances
NJ Tree

Cladograms

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

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
4 5 3 4 627 12 3 0 
                 +-- 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
3 3 1 3 26 14 3 2 
  +----------------------- 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
4 4 2 4 23 13 3 3 
  +----------------- Theliderma        
--+                                    
  |  +-------------- Cyclonaias        
  +--+                                 
     |  +----------- Pustulosa         
     +--+                              
        |  +-------- Quadrula_nobilis **  
        +--+                           
           |  +----- Quadrula_verrucosa **
           +--+                        
              |  +-- Quadrula_fragosa **  
              +--+                     
                 +-- Quadrula_quadrula **

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

iTtTbiriSoTroTroSiD
3 9 6 3 70 6 3 7 
              +-- 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 11 
  +----------------------- 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 7 
  +-------------- Theliderma          
  |                                   
--+  +----------- Quadrula_quadrula **   
  |  |                                
  |  |        +-- Quadrula_nobilis **    
  +--+  +-----+                       
     |  |     +-- Quadrula_verrucosa **  
     |  |                             
     +--+  +----- Pustulosa_kieneriana
        |  |                          
        |  |  +-- Cyclonaias          
        +--+--+                       
           |  +-- Pustulosa_succissa  
           |                          
           |  +-- Pustulosa_infucata  
           +--+                       
              +-- Pustulosa_kleiniana

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

iTtTbiriSoTroTroSiD
3 13 11 3 26 10 2 10 
        +-- 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

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

iTtTbiriSoTroTroSiD
3 3 2 4 11 10 2 5 
  +----------- Pustulosa           
--+                                
  |  +-------- Quadrula_quadrula__1 **
  +--+                             
     |  +----- Quadrula_verrucosa **  
     +--+                          
        |  +-- Quadrula_nobilis **    
        +--+                       
           +-- Quadrula_quadrula__2 **

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

iTtTbiriSoTroTroSiD
3 8 6 3 70 6 3 9 
  +----------------- Theliderma         
  |                                     
--+  +-------------- Quadrula_verrucosa ** 
  |  |                                  
  |  |           +-- Quadrula_quadrula **  
  +--+  +--------+                      
     |  |        +-- Quadrula_nobilis **   
     |  |                               
     +--+        +-- Pustulosa_infucata 
        |  +-----+                      
        |  |     +-- Pustulosa_kleiniana
        +--+                            
           |  +----- Cyclonaias         
           +--+                         
              |  +-- Pustulosa_pustulosa
              +--+                      
                 +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
3 3 1 3 26 10 2 4 
  +----------- Theliderma        
--+                              
  |  +-------- Pustulosa         
  +--+                           
     |  +----- Quadrula_verrucosa **
     +--+                        
        |  +-- Quadrula_quadrula ** 
        +--+                     
           +-- Quadrula_nobilis **

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

iTtTbiriSoTroTroSiD
3 3 1 3 99 6 3 4 
  +----------- Theliderma          
  |                                
  |     +----- Quadrula_verrucosa **  
--+  +--+                          
  |  |  |  +-- Quadrula_quadrula **   
  |  |  +--+                       
  |  |     +-- Quadrula_nobilis **    
  +--+                             
     |  +----- Pustulosa_kieneriana
     |  |                          
     |  |  +-- Cyclonaias          
     +--+--+                       
        |  +-- Pustulosa_succissa  
        |                          
        |  +-- Pustulosa_infucata  
        +--+                       
           +-- Pustulosa_kleiniana

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

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

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

iTtTbiriSoTroTroSiD
2 5 0 2 103 3 3 7 
  +-- Cyclonaias            
  |                         
  +-- Pustulosa_pustulosa__1
  |                         
  +-- Theliderma            
--+                         
  +-- Pustulosa_pustulosa__2
  |                         
  +-- Quadrula_quadrula **     
  |                         
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 198 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 136 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 9 0 2 122 0 0 10 
  +-- Amblema           
  |                     
  +-- Ortmanniana       
  |                     
  +-- Lampsilis         
  |                     
  +-- Cambarunio        
  |                     
--+-- Elliptio          
  |                     
  +-- Eurynia           
  |                     
  +-- Pleurobema        
  |                     
  +-- Quadrula_quadrula ** 
  |                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 5 0 2 36 3 2 6 
  +----------- 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 6 
  +-------------- Theliderma          
--+                                   
  |  +----------- Quadrula_quadrula **   
  +--+                                
     |  +-------- Pustulosa_infucata  
     +--+                             
        |  +----- Quadrula_verrucosa **  
        +--+                          
           |  +-- Pustulosa_kieneriana
           +--+                       
              +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
2 2 0 2 196 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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 8 
  +-------------- Quadrula_quadrula **  
--+                                  
  |  +----------- Quadrula_verrucosa ** 
  +--+                               
     |  +-------- Theliderma         
     +--+                            
        |  +----- Pustulosa_pustulosa
        +--+                         
           |  +-- Cyclonaias         
           +--+                      
              +-- Pustulosa_nodulata

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

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

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

iTtTbiriSoTroTroSiD
2 2 0 2 198 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 **

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
2 15 0 2 13 0 0 17 
  +-------------------- 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. S1

iTtTbiriSoTroTroSiD
2 2 0 2 136 0 0 -- 
  +-- Quadrula_quadrula ** 
--+                     
  +-- Quadrula_verrucosa **

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

iTtTbiriSoTroTroSiD
2 3 0 2 29 1 1 4 
  +----- Quadrula_verrucosa **
--+                        
  |  +-- Cyclonaias        
  +--+                     
     +-- Quadrula_quadrula **

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 **

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 22 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., 2023 [Publ 4339], fig. 5

iTtTbiriSoTroTroSiD
1 1 0 1 18 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 **

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 140 0 0 -- 
-- Quadrula_quadrula **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 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

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

iTtTbiriSoTroTroSiD
1 1 0 1 81 0 0 -- 
-- Quadrula_quadrula **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 91 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. 3

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Krebs, 2004 [Publ 2692], fig. 2

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 37 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. 4

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 30 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. 2c

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 5 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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 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., 2021 [Publ 4366], fig. 3-ML

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 0 0 -- 
-- Quadrula_quadrula **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 **

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

iTtTbiriSoTroTroSiD
1 1 0 1 37 2 2 -- 
  +----- Quadrula_quadrula **
--+                       
  |  +-- Cyclonaias       
  +--+                    
     +-- Pustulosa

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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 **

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

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

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 **