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Page last updated
Thu 17 July 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!

Bivalvia | ... | Unionoidea | Unionidae | Ambleminae | Quadrulini | Quadrulina

genus Pustulosa Frierson, 1927: cladomics
     9 Recent valid species (10 total species)

Cladomics Summary

number of species = 9 (10 total)
analyzed in 41 cladograms in 23 publications
maximum number of terminals in an analysis = 8 (mean = 3.02, median = 2)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 24
maximum number of included terminals = 8, (mean = 4.46 median = 4)
proportion of cladograms that support basic monophyly = 0.25

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

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

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

iTtTbiriSoTroTroSiD
897821930
  +----------------------- Theliderma          
  |                                            
--+  +-------------------- Quadrula            
  |  |                                         
  +--+  +----------------- Pustulosa_kieneriana **
     |  |                                      
     |  |              +-- Pustulosa_infucata **  
     +--+  +-----------+                       
        |  |           +-- Pustulosa_kleiniana ** 
        |  |                                   
        +--+           +-- Pustulosa_pustulosa ** 
           |  +--------+                       
           |  |        +-- Pustulosa_succissa **  
           +--+                                
              |  +-------- Cyclonaias          
              +--+                             
                 |  +----- Pustulosa_necki **     
                 +--+                          
                    |  +-- Pustulosa_petrina **   
                    +--+                       
                       +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
897820636
  +----------------------- Quadrula_quadrula   
  |                                            
  |                    +-- Quadrula_verrucosa  
--+  +-----------------+                       
  |  |                 +-- Quadrula_nobilis    
  |  |                                         
  +--+  +----------------- Theliderma          
     |  |                                      
     |  |           +----- Pustulosa_kieneriana **
     +--+  +--------+                          
        |  |        |  +-- Pustulosa_infucata **  
        |  |        +--+                       
        +--+           +-- Pustulosa_kleiniana ** 
           |                                   
           |  +----------- Cyclonaias          
           |  |                                
           +--+        +-- Pustulosa_pustulosa ** 
              |  +-----+                       
              |  |     +-- Pustulosa_succissa **  
              +--+                             
                 |  +----- Pustulosa_nodulata **  
                 +--+                          
                    |  +-- Pustulosa_necki **     
                    +--+                       
                       +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
886819939
  +-------------------- Theliderma          
  |                                         
--+  +----------------- Cyclonaias          
  |  |                                      
  +--+  +-------------- Quadrula            
     |  |                                   
     |  |        +----- Pustulosa_nodulata **  
     +--+  +-----+                          
        |  |     |  +-- Pustulosa_necki **     
        |  |     +--+                       
        |  |        +-- Pustulosa_petrina **   
        +--+                                
           |        +-- Pustulosa_kieneriana **
           |  +-----+                       
           |  |     +-- Pustulosa_pustulosa ** 
           +--+                             
              |  +----- Pustulosa_succissa **  
              +--+                          
                 |  +-- Pustulosa_infucata **  
                 +--+                       
                    +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
897818632
  +----------------------- Theliderma          
  |                                            
--+  +-------------------- Quadrula_quadrula   
  |  |                                         
  |  |                 +-- Quadrula_verrucosa  
  +--+  +--------------+                       
     |  |              +-- Quadrula_nobilis    
     |  |                                      
     |  |           +----- Pustulosa_kieneriana **
     +--+  +--------+                          
        |  |        |  +-- Pustulosa_infucata **  
        |  |        +--+                       
        |  |           +-- Pustulosa_kleiniana ** 
        +--+                                   
           |           +-- Pustulosa_pustulosa ** 
           |  +--------+                       
           |  |        +-- Pustulosa_succissa **  
           +--+                                
              |  +-------- Cyclonaias          
              +--+                             
                 |  +----- Pustulosa_necki **     
                 +--+                          
                    |  +-- Pustulosa_petrina **   
                    +--+                       
                       +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
89788432
  +-------------------- Theliderma          
  |                                         
--+  +----------------- Quadrula            
  |  |                                      
  |  |           +----- Pustulosa_kieneriana **
  +--+  +--------+                          
     |  |        |  +-- Pustulosa_infucata **  
     |  |        +--+                       
     |  |           +-- Pustulosa_kleiniana ** 
     +--+                                   
        |           +-- Pustulosa_pustulosa ** 
        |  +--------+                       
        |  |        +-- Pustulosa_succissa **  
        +--+                                
           |  +-------- Cyclonaias          
           +--+                             
              |  +----- Pustulosa_necki **     
              +--+                          
                 |  +-- Pustulosa_nodulata **  
                 +--+                       
                    +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
663623724
  +-------------- Theliderma          
  |                                   
--+  +----------- Quadrula            
  |  |                                
  +--+  +-------- Pustulosa_kieneriana **
     |  |                             
     |  +-------- Pustulosa_infucata **  
     +--+                             
        |     +-- Pustulosa_pustulosa ** 
        |  +--+                       
        |  |  +-- Pustulosa_succissa **  
        +--+                          
           |  +-- Pustulosa_nodulata **  
           +--+                       
              +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
664623723
                    +-- Quadrula_verrucosa  
  +-----------------+                       
  |                 +-- Quadrula_nobilis    
--+                                         
  |  +----------------- Quadrula_quadrula   
  |  |                                      
  +--+  +-------------- Theliderma          
     |  |                                   
     +--+  +----------- Pustulosa_kieneriana **
        |  |                                
        +--+  +-------- Pustulosa_infucata **  
           |  |                             
           +--+     +-- Pustulosa_succissa **  
              |  +--+                       
              |  |  +-- Pustulosa_pustulosa ** 
              +--+                          
                 |  +-- Pustulosa_nodulata **  
                 +--+                       
                    +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
66468723
  +-------------- Quadrula            
  |                                   
--+  +----------- Pustulosa_kieneriana **
  |  |                                
  +--+  +-------- Pustulosa_infucata **  
     |  |                             
     +--+     +-- Pustulosa_pustulosa ** 
        |  +--+                       
        |  |  +-- Pustulosa_succissa **  
        +--+                          
           |  +-- Pustulosa_nodulata **  
           +--+                       
              +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
453469534
  +----------------- Theliderma         
  |                                     
--+  +-------------- Quadrula_verrucosa 
  |  |                                  
  |  |           +-- Quadrula_quadrula  
  +--+  +--------+                      
     |  |        +-- Quadrula_nobilis   
     |  |                               
     +--+        +-- Pustulosa_infucata ** 
        |  +-----+                      
        |  |     +-- Pustulosa_kleiniana **
        +--+                            
           |  +----- Cyclonaias         
           +--+                         
              |  +-- Pustulosa_pustulosa **
              +--+                      
                 +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
453438224
  +----------- Quadrula           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
453438224
  +----------- Quadrula           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
464439225
           +-- Pustulosa_infucata ** 
  +--------+                      
  |        +-- Pustulosa_kleiniana **
--+                               
  |  +-------- Quadrula           
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
453438224
  +----------- Quadrula           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
452498535
  +-------------- Theliderma          
  |                                   
--+  +----------- Quadrula_quadrula   
  |  |                                
  |  |        +-- Quadrula_nobilis    
  +--+  +-----+                       
     |  |     +-- Quadrula_verrucosa  
     |  |                             
     +--+  +----- Pustulosa_kieneriana **
        |  |                          
        |  |  +-- Cyclonaias          
        +--+--+                       
           |  +-- Pustulosa_succissa **  
           |                          
           |  +-- Pustulosa_infucata **  
           +--+                       
              +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
453469534
              +-- Quadrula_verrucosa 
  +-----------+                      
  |           +-- Quadrula_nobilis   
  |                                  
--+  +----------- Quadrula_quadrula  
  |  |                               
  |  +----------- Theliderma         
  +--+                               
     |        +-- Pustulosa_infucata ** 
     |  +-----+                      
     |  |     +-- Pustulosa_kleiniana **
     +--+                            
        |  +----- Cyclonaias         
        +--+                         
           |  +-- Pustulosa_pustulosa **
           +--+                      
              +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
452498535
  +----------- Theliderma          
  |                                
--+  +-------- Quadrula            
  |  |                             
  +--+  +----- Pustulosa_kieneriana **
     |  |                          
     |  |  +-- Cyclonaias          
     +--+--+                       
        |  +-- Pustulosa_succissa **  
        |                          
        |  +-- Pustulosa_infucata **  
        +--+                       
           +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
342315429
  +-------------- Theliderma          
--+                                   
  |  +----------- Quadrula_quadrula   
  +--+                                
     |  +-------- Pustulosa_infucata **  
     +--+                             
        |  +----- Quadrula_verrucosa  
        +--+                          
           |  +-- Pustulosa_kieneriana **
           +--+                       
              +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
230212118
  +-- Quadrula          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
230236436
  +----------- Theliderma         
--+                               
  |  +-------- Quadrula           
  +--+                            
     |  +----- Pustulosa_pustulosa **
     +--+                         
        |  +-- Cyclonaias         
        +--+                      
           +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
230236326
  +----------- Quadrula_quadrula  
--+                               
  |  +-------- Quadrula_verrucosa 
  +--+                            
     |  +----- Pustulosa_pustulosa **
     +--+                         
        |  +-- Pustulosa_petrina **  
        +--+                      
           +-- Cyclonaias

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

iTtTbiriSoTroTroSiD
230233436
  +-------------- Quadrula_quadrula  
--+                                  
  |  +----------- Quadrula_verrucosa 
  +--+                               
     |  +-------- Theliderma         
     +--+                            
        |  +----- Pustulosa_pustulosa **
        +--+                         
           |  +-- Cyclonaias         
           +--+                      
              +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
22021211--
  +----- Quadrula          
--+                        
  |  +-- Pustulosa_infucata **
  +--+                     
     +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
230212118
  +-- Quadrula          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
22021211--
  +----- Quadrula          
--+                        
  |  +-- Pustulosa_infucata **
  +--+                     
     +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
15021034311
  +-- Cyclonaias            
  |                         
  +-- Pustulosa_pustulosa__1 **
  |                         
  +-- Theliderma            
--+                         
  +-- Pustulosa_pustulosa__2 **
  |                         
  +-- Quadrula_quadrula     
  |                         
  +-- Quadrula_verrucosa

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

iTtTbiriSoTroTroSiD
1101311--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11013711--
  +-- Cyclonaias         
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11013622--
  +----- Quadrula           
--+                         
  |  +-- Cyclonaias         
  +--+                      
     +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1101311--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11011111--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11011911--
  +-- Quadrula         
--+                    
  +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
11014011--
  +-- Pustulosa_petrina **
--+                    
  +-- Quadrula

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

iTtTbiriSoTroTroSiD
11011011--
-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11012011--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11012011--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11011011--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1101311--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11011111--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11011011--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11011111--
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
11014111--
  +-- Pustulosa_petrina **
--+                    
  +-- Quadrula
 
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