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

genus Pustulosa Frierson, 1927: cladomics

8 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 = 8 (9 total)
analyzed in 42 cladograms in 24 publications
maximum number of terminals in an analysis = 8 (mean = 3.14, median = 2)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 25
maximum number of included terminals = 8, (mean = 4.6 median = 4)
proportion of cladograms that support basic monophyly = 0.24

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 Pustulosa cladogram distances
PCoA Ordination

NJ Tree of Pustulosa cladogram distances
NJ Tree

Cladograms

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

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

iTtTbiriSoTroTroSiD
8 9 7 8 21 9 3 0 
  +----------------------- 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. 4

iTtTbiriSoTroTroSiD
8 9 7 8 18 6 3 2 
  +----------------------- Theliderma          
  |                                            
--+  +-------------------- Quadrula_quadrula   
  |  |                                         
  |  |                 +-- Quadrula_verrucosa  
  +--+  +--------------+                       
     |  |              +-- Quadrula_nobilis    
     |  |                                      
     |  |           +----- 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
8 8 6 8 19 9 3 9 
  +-------------------- Theliderma          
  |                                         
--+  +----------------- Cyclonaias          
  |  |                                      
  +--+  +-------------- Quadrula            
     |  |                                   
     |  |        +----- Pustulosa_nodulata **  
     +--+  +-----+                          
        |  |     |  +-- Pustulosa_necki **     
        |  |     +--+                       
        |  |        +-- Pustulosa_petrina **   
        +--+                                
           |        +-- Pustulosa_kieneriana **
           |  +-----+                       
           |  |     +-- Pustulosa_pustulosa ** 
           +--+                             
              |  +----- Pustulosa_succissa **  
              +--+                          
                 |  +-- Pustulosa_infucata **  
                 +--+                       
                    +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
8 9 7 8 8 4 3 2 
  +-------------------- Theliderma          
  |                                         
--+  +----------------- Quadrula            
  |  |                                      
  |  |           +----- Pustulosa_kieneriana **
  +--+  +--------+                          
     |  |        |  +-- Pustulosa_infucata **  
     |  |        +--+                       
     |  |           +-- Pustulosa_kleiniana ** 
     +--+                                   
        |           +-- Pustulosa_pustulosa ** 
        |  +--------+                       
        |  |        +-- Pustulosa_succissa **  
        +--+                                
           |  +-------- Cyclonaias          
           +--+                             
              |  +----- Pustulosa_necki **     
              +--+                          
                 |  +-- Pustulosa_nodulata **  
                 +--+                       
                    +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
8 9 7 8 20 6 3 6 
  +----------------------- Quadrula_quadrula   
  |                                            
  |                    +-- Quadrula_verrucosa  
--+  +-----------------+                       
  |  |                 +-- Quadrula_nobilis    
  |  |                                         
  +--+  +----------------- Theliderma          
     |  |                                      
     |  |           +----- Pustulosa_kieneriana **
     +--+  +--------+                          
        |  |        |  +-- Pustulosa_infucata **  
        |  |        +--+                       
        +--+           +-- Pustulosa_kleiniana ** 
           |                                   
           |  +----------- Cyclonaias          
           |  |                                
           +--+        +-- Pustulosa_pustulosa ** 
              |  +-----+                       
              |  |     +-- Pustulosa_succissa **  
              +--+                             
                 |  +----- Pustulosa_nodulata **  
                 +--+                          
                    |  +-- Pustulosa_necki **     
                    +--+                       
                       +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
8 9 7 8 623 8 3 6 
                 +-- 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 **

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

iTtTbiriSoTroTroSiD
6 6 4 6 23 7 2 3 
                    +-- 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
6 6 4 6 8 7 2 3 
  +-------------- Quadrula            
  |                                   
--+  +----------- Pustulosa_kieneriana **
  |  |                                
  +--+  +-------- Pustulosa_infucata **  
     |  |                             
     +--+     +-- Pustulosa_pustulosa ** 
        |  +--+                       
        |  |  +-- Pustulosa_succissa **  
        +--+                          
           |  +-- Pustulosa_nodulata **  
           +--+                       
              +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
6 6 3 6 23 7 2 4 
  +-------------- Theliderma          
  |                                   
--+  +----------- Quadrula            
  |  |                                
  +--+  +-------- Pustulosa_kieneriana **
     |  |                             
     |  +-------- Pustulosa_infucata **  
     +--+                             
        |     +-- Pustulosa_pustulosa ** 
        |  +--+                       
        |  |  +-- Pustulosa_succissa **  
        +--+                          
           |  +-- Pustulosa_nodulata **  
           +--+                       
              +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
4 5 3 4 38 2 2 4 
  +----------- Quadrula           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
4 5 3 4 38 2 2 4 
  +----------- Quadrula           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
4 5 2 4 98 5 3 5 
  +----------- Theliderma          
  |                                
--+  +-------- Quadrula            
  |  |                             
  +--+  +----- Pustulosa_kieneriana **
     |  |                          
     |  |  +-- Cyclonaias          
     +--+--+                       
        |  +-- Pustulosa_succissa **  
        |                          
        |  +-- Pustulosa_infucata **  
        +--+                       
           +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
4 5 2 4 98 5 3 5 
  +-------------- Theliderma          
  |                                   
--+  +----------- Quadrula_quadrula   
  |  |                                
  |  |        +-- Quadrula_nobilis    
  +--+  +-----+                       
     |  |     +-- Quadrula_verrucosa  
     |  |                             
     +--+  +----- Pustulosa_kieneriana **
        |  |                          
        |  |  +-- Cyclonaias          
        +--+--+                       
           |  +-- Pustulosa_succissa **  
           |                          
           |  +-- Pustulosa_infucata **  
           +--+                       
              +-- Pustulosa_kleiniana **

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

iTtTbiriSoTroTroSiD
4 6 4 4 39 2 2 5 
           +-- Pustulosa_infucata ** 
  +--------+                      
  |        +-- Pustulosa_kleiniana **
--+                               
  |  +-------- Quadrula           
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
4 5 3 4 38 2 2 4 
  +----------- Quadrula           
  |                               
--+        +-- Pustulosa_infucata ** 
  |  +-----+                      
  |  |     +-- Pustulosa_kleiniana **
  +--+                            
     |  +----- Cyclonaias         
     +--+                         
        |  +-- Pustulosa_succissa ** 
        +--+                      
           +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
4 5 3 4 69 5 3 4 
  +----------------- Theliderma         
  |                                     
--+  +-------------- Quadrula_verrucosa 
  |  |                                  
  |  |           +-- Quadrula_quadrula  
  +--+  +--------+                      
     |  |        +-- Quadrula_nobilis   
     |  |                               
     +--+        +-- Pustulosa_infucata ** 
        |  +-----+                      
        |  |     +-- Pustulosa_kleiniana **
        +--+                            
           |  +----- Cyclonaias         
           +--+                         
              |  +-- Pustulosa_pustulosa **
              +--+                      
                 +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
4 5 3 4 69 5 3 4 
              +-- Quadrula_verrucosa 
  +-----------+                      
  |           +-- Quadrula_nobilis   
  |                                  
--+  +----------- Quadrula_quadrula  
  |  |                               
  |  +----------- Theliderma         
  +--+                               
     |        +-- Pustulosa_infucata ** 
     |  +-----+                      
     |  |     +-- Pustulosa_kleiniana **
     +--+                            
        |  +----- Cyclonaias         
        +--+                         
           |  +-- Pustulosa_pustulosa **
           +--+                      
              +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
3 4 2 3 15 4 2 9 
  +-------------- Theliderma          
--+                                   
  |  +----------- Quadrula_quadrula   
  +--+                                
     |  +-------- Pustulosa_infucata **  
     +--+                             
        |  +----- Quadrula_verrucosa  
        +--+                          
           |  +-- Pustulosa_kieneriana **
           +--+                       
              +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
2 2 0 2 12 1 1 -- 
  +----- Quadrula          
--+                        
  |  +-- Pustulosa_infucata **
  +--+                     
     +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
2 3 0 2 36 3 2 6 
  +----------- Quadrula_quadrula  
--+                               
  |  +-------- Quadrula_verrucosa 
  +--+                            
     |  +----- Pustulosa_pustulosa **
     +--+                         
        |  +-- Pustulosa_petrina **  
        +--+                      
           +-- Cyclonaias

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

iTtTbiriSoTroTroSiD
2 2 0 2 12 1 1 -- 
  +----- Quadrula          
--+                        
  |  +-- Pustulosa_infucata **
  +--+                     
     +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
2 3 0 2 12 1 1 8 
  +-- Quadrula          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

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

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

iTtTbiriSoTroTroSiD
2 3 0 2 12 1 1 8 
  +-- Quadrula          
  |                     
--+-- Pustulosa_infucata **
  |                     
  +-- Pustulosa_succissa **

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

iTtTbiriSoTroTroSiD
2 3 0 2 36 4 3 6 
  +----------- Theliderma         
--+                               
  |  +-------- Quadrula           
  +--+                            
     |  +----- Pustulosa_pustulosa **
     +--+                         
        |  +-- Cyclonaias         
        +--+                      
           +-- Pustulosa_nodulata **

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Pustulosa_petrina **
--+                    
  +-- Quadrula

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

iTtTbiriSoTroTroSiD
1 5 0 2 104 4 3 11 
  +-- Cyclonaias            
  |                         
  +-- Pustulosa_pustulosa__1 **
  |                         
  +-- Theliderma            
--+                         
  +-- Pustulosa_pustulosa__2 **
  |                         
  +-- Quadrula_quadrula     
  |                         
  +-- Quadrula_verrucosa

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 1 1 -- 
-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 20 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 37 2 2 -- 
  +----- Quadrula           
--+                         
  |  +-- Cyclonaias         
  +--+                      
     +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 19 1 1 -- 
  +-- Quadrula         
--+                    
  +-- Pustulosa_petrina **

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

iTtTbiriSoTroTroSiD
1 1 0 1 38 1 1 -- 
  +-- Cyclonaias         
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 11 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 1 1 -- 
  +-- Pustulosa_petrina **
--+                    
  +-- Quadrula

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

iTtTbiriSoTroTroSiD
1 1 0 1 3 1 1 -- 
  +-- Quadrula           
--+                      
  +-- Pustulosa_pustulosa **