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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

genus Uniomerus Conrad, 1853: cladomics
     4 Recent valid species (5 total species)

Cladomics Summary

number of species = 4 (5 total)
analyzed in 51 cladograms in 35 publications
maximum number of terminals in an analysis = 4 (mean = 1.2, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 6
maximum number of included terminals = 4, (mean = 2.67 median = 2.5)
proportion of cladograms that support basic monophyly = 1

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 ** ** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
33246291820
           +----------- Megalonaias           
  +--------+                                  
  |        |  +-------- Uniomerus_carolinianus **
  |        +--+                               
  |           |  +----- Uniomerus_declivis__2 ** 
  |           +--+                            
  |              |  +-- Uniomerus_declivis__1 ** 
  |              +--+                         
  |                 +-- Uniomerus_tetralasmus ** 
  |                                           
  |                 +-- 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

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

iTtTbiriSoTroTroSiD
4424231822
  +-------------- Megalonaias           
  |                                     
  |     +-------- Uniomerus_carolinianus **
--+  +--+                               
  |  |  |  +----- Uniomerus_columbensis ** 
  |  |  +--+                            
  |  |     |  +-- Uniomerus_tetralasmus ** 
  +--+     +--+                         
     |        +-- Uniomerus_declivis **    
     |                                  
     |  +-------- Theliderma            
     +--+                               
        |  +----- Cyclonaias            
        +--+                            
           |  +-- Quadrula              
           +--+                         
              +-- Pustulosa

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

iTtTbiriSoTroTroSiD
3303251522
                       +----- Megalonaias           
                       |                            
  +--------------------+  +-- Uniomerus_carolinianus **
  |                    |  |                         
  |                    +--+-- Uniomerus_declivis **    
  |                       |                         
--+                       +-- Uniomerus_tetralasmus ** 
  |                                                 
  |  +----------------------- 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, 2012 [Publ 3309], fig. 3

iTtTbiriSoTroTroSiD
22027191--
                 +-- Uniomerus_declivis **   
  +--------------+                        
  |              +-- Uniomerus_tetralasmus **
  |                                       
--+              +-- Quadrula_verrucosa   
  |  +-----------+                        
  |  |           +-- Quadrula_nobilis     
  |  |                                    
  +--+  +----------- Quadrula_quadrula    
     |  |                                 
     |  +----------- Theliderma           
     +--+                                 
        |        +-- Pustulosa_infucata   
        |  +-----+                        
        |  |     +-- Pustulosa_kleiniana  
        +--+                              
           |  +----- Cyclonaias           
           +--+                           
              |  +-- Pustulosa_pustulosa  
              +--+                        
                 +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
22027191--
                    +-- Uniomerus_declivis **   
  +-----------------+                        
  |                 +-- Uniomerus_tetralasmus **
--+                                          
  |  +----------------- Theliderma           
  |  |                                       
  +--+  +-------------- Quadrula_verrucosa   
     |  |                                    
     |  |           +-- Quadrula_quadrula    
     +--+  +--------+                        
        |  |        +-- Quadrula_nobilis     
        |  |                                 
        +--+        +-- Pustulosa_infucata   
           |  +-----+                        
           |  |     +-- Pustulosa_kleiniana  
           +--+                              
              |  +----- Cyclonaias           
              +--+                           
                 |  +-- Pustulosa_pustulosa  
                 +--+                        
                    +-- Pustulosa_succissa

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

iTtTbiriSoTroTroSiD
220210351--
     +-- Uniomerus_declivis **    
  +--+                         
  |  +-- Uniomerus_tetralasmus ** 
  |                            
  |  +-- Cyclonaias            
--+  |                         
  |  +-- Pustulosa_pustulosa__1
  |  |                         
  |  +-- Theliderma            
  +--+                         
     +-- Pustulosa_pustulosa__2
     |                         
     +-- Quadrula_quadrula     
     |                         
     +-- Quadrula_verrucosa

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

iTtTbiriSoTroTroSiD
11013042--
  +----------- Megalonaias       
--+                              
  |  +-------- Uniomerus_declivis **
  +--+                           
     |  +----- Quadrula_verrucosa
     +--+                        
        |  +-- Cyclonaias        
        +--+                     
           +-- Quadrula_quadrula

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

iTtTbiriSoTroTroSiD
11013211--
  +-- Quadrula          
--+                     
  +-- Uniomerus_declivis **

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

iTtTbiriSoTroTroSiD
11019511--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11012132--
-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
110119932--
-- Uniomerus_carolinianus **

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

iTtTbiriSoTroTroSiD
110128182--
  +-------------------------- Uniomerus_tetralasmus **
  |                                                
--+  +----------------------- Theliderma           
  |  |                                             
  +--+  +-------------------- Quadrula             
     |  |                                          
     +--+  +----------------- Pustulosa_kieneriana 
        |  |                                       
        |  |              +-- Pustulosa_infucata   
        +--+  +-----------+                        
           |  |           +-- Pustulosa_kleiniana  
           |  |                                    
           +--+           +-- Pustulosa_pustulosa  
              |  +--------+                        
              |  |        +-- Pustulosa_succissa   
              +--+                                 
                 |  +-------- Cyclonaias           
                 +--+                              
                    |  +----- Pustulosa_necki      
                    +--+                           
                       |  +-- Pustulosa_petrina    
                       +--+                        
                          +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11018711--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11012732--
-- Uniomerus_carolinianus **

Campbell et al., 2008 [Publ 2675], fig. 5

iTtTbiriSoTroTroSiD
11012400--
-- Uniomerus_declivis **

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

iTtTbiriSoTroTroSiD
110115132--
  +-------------------------- Megalonaias          
  |                                                
--+  +----------------------- Uniomerus_tetralasmus **
  |  |                                             
  +--+  +-------------------- Theliderma           
     |  |                                          
     +--+  +----------------- Quadrula             
        |  |                                       
        |  |           +----- Pustulosa_kieneriana 
        +--+  +--------+                           
           |  |        |  +-- Pustulosa_infucata   
           |  |        +--+                        
           |  |           +-- Pustulosa_kleiniana  
           +--+                                    
              |           +-- Pustulosa_pustulosa  
              |  +--------+                        
              |  |        +-- Pustulosa_succissa   
              +--+                                 
                 |  +-------- Cyclonaias           
                 +--+                              
                    |  +----- Pustulosa_necki      
                    +--+                           
                       |  +-- Pustulosa_nodulata   
                       +--+                        
                          +-- Pustulosa_petrina

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

iTtTbiriSoTroTroSiD
11011832--
-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
1101332--
  +----- Quadrula          
--+                        
  |  +-- Uniomerus_declivis **
  +--+                     
     +-- Megalonaias

Huang et al., 2002 [Publ 2657], fig. 1

iTtTbiriSoTroTroSiD
11013311--
-- Uniomerus_carolinianus **

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

iTtTbiriSoTroTroSiD
1101101102--
  +-------------------- Megalonaias         
  |                                         
--+  +----------------- Uniomerus_declivis **  
  |  |                                      
  +--+  +-------------- Theliderma          
     |  |                                   
     +--+  +----------- Quadrula_quadrula   
        |  |                                
        |  |        +-- Quadrula_nobilis    
        +--+  +-----+                       
           |  |     +-- Quadrula_verrucosa  
           |  |                             
           +--+  +----- Pustulosa_kieneriana
              |  |                          
              |  |  +-- Cyclonaias          
              +--+--+                       
                 |  +-- Pustulosa_succissa  
                 |                          
                 |  +-- Pustulosa_infucata  
                 +--+                       
                    +-- Pustulosa_kleiniana

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

iTtTbiriSoTroTroSiD
11015511--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11015611--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

Kandl et al., 2001 [Publ 3384], fig. 4

iTtTbiriSoTroTroSiD
1101500--
-- Uniomerus_carolinianus **

Campbell et al., 2008 [Publ 2675], fig. 4

iTtTbiriSoTroTroSiD
11012400--
-- Uniomerus_declivis **

Perkins et al., 2017 [Publ 3777], fig. 3

iTtTbiriSoTroTroSiD
11012300--
-- Uniomerus_declivis **

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

iTtTbiriSoTroTroSiD
11019111--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11011811--
  +-- Uniomerus_tetralasmus **
--+                        
  +-- Quadrula

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

iTtTbiriSoTroTroSiD
11019511--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
110125152--
  +----------------------------- Megalonaias          
  |                                                   
--+  +-------------------------- Uniomerus_tetralasmus **
  |  |                                                
  +--+  +----------------------- Theliderma           
     |  |                                             
     +--+  +-------------------- Quadrula_quadrula    
        |  |                                          
        |  |                 +-- Quadrula_verrucosa   
        +--+  +--------------+                        
           |  |              +-- Quadrula_nobilis     
           |  |                                       
           |  |           +----- Pustulosa_kieneriana 
           +--+  +--------+                           
              |  |        |  +-- Pustulosa_infucata   
              |  |        +--+                        
              |  |           +-- Pustulosa_kleiniana  
              +--+                                    
                 |           +-- Pustulosa_pustulosa  
                 |  +--------+                        
                 |  |        +-- Pustulosa_succissa   
                 +--+                                 
                    |  +-------- Cyclonaias           
                    +--+                              
                       |  +----- Pustulosa_necki      
                       +--+                           
                          |  +-- Pustulosa_petrina    
                          +--+                        
                             +-- Pustulosa_nodulata

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

iTtTbiriSoTroTroSiD
11017311--
  +-- Uniomerus_tetralasmus **
--+                        
  +-- Quadrula

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

iTtTbiriSoTroTroSiD
11017311--
  +-- Uniomerus_tetralasmus **
--+                        
  +-- Quadrula

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

iTtTbiriSoTroTroSiD
110119732--
  +----- Quadrula              
--+                            
  |  +-- Megalonaias           
  +--+                         
     +-- Uniomerus_carolinianus **

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

iTtTbiriSoTroTroSiD
110119932--
  +----- Quadrula              
--+                            
  |  +-- Megalonaias           
  +--+                         
     +-- Uniomerus_carolinianus **

Huang et al., 2002 [Publ 2657], fig. 2

iTtTbiriSoTroTroSiD
11013311--
-- Uniomerus_carolinianus **

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

iTtTbiriSoTroTroSiD
11018511--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11015611--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11014261--
  +-------------- Uniomerus_declivis ** 
  |                                  
--+           +-- Pustulosa_infucata 
  |  +--------+                      
  |  |        +-- Pustulosa_kleiniana
  +--+                               
     |  +-------- Quadrula           
     +--+                            
        |  +----- Cyclonaias         
        +--+                         
           |  +-- Pustulosa_succissa 
           +--+                      
              +-- Pustulosa_pustulosa

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

iTtTbiriSoTroTroSiD
11012132--
-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
110114011--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11011411--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11013232--
-- Uniomerus_carolinianus **

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

iTtTbiriSoTroTroSiD
11012632--
-- Uniomerus_carolinianus **

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

iTtTbiriSoTroTroSiD
11019111--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11013751--
  +-------------- Uniomerus_declivis ** 
--+                                  
  |  +----------- Quadrula_quadrula  
  +--+                               
     |  +-------- Quadrula_verrucosa 
     +--+                            
        |  +----- Pustulosa_pustulosa
        +--+                         
           |  +-- Pustulosa_petrina  
           +--+                      
              +-- Cyclonaias

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
11011911--
  +-- Uniomerus_tetralasmus **
--+                        
  +-- Megalonaias

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11015311--
  +-- Quadrula             
--+                        
  +-- Uniomerus_tetralasmus **

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

iTtTbiriSoTroTroSiD
11017311--
  +-- Uniomerus_tetralasmus **
--+                        
  +-- Quadrula

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

iTtTbiriSoTroTroSiD
1101101102--
              +-- Megalonaias         
  +-----------+                       
  |           +-- Uniomerus_declivis **  
--+                                   
  |  +----------- Theliderma          
  |  |                                
  +--+  +-------- Quadrula            
     |  |                             
     +--+  +----- Pustulosa_kieneriana
        |  |                          
        |  |  +-- Cyclonaias          
        +--+--+                       
           |  +-- Pustulosa_succissa  
           |                          
           |  +-- Pustulosa_infucata  
           +--+                       
              +-- Pustulosa_kleiniana
 
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