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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 | Margaritiferidae | Margaritiferinae

genus Pseudunio Haas, 1910: cladomics
     3 Recent valid species (5 total species)

Cladomics Summary

number of species = 3 (5 total)
analyzed in 81 cladograms in 44 publications
maximum number of terminals in an analysis = 5 (mean = 1.53, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 31
maximum number of included terminals = 5, (mean = 2.39 median = 2)
proportion of cladograms that support basic monophyly = 0.94

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

Lyubas et al., 2019 [Publ 3963], fig. 10

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

iTtTbiriSoTroTroSiD
551511820
  +----------- Margaritifera             
  |                                      
--+  +-------- Cumberlandia              
  |  |                                   
  +--+  +----- Pseudunio_marocanus **       
     |  |                                
     +--+  +-- Pseudunio_auricularius **    
        |  |                             
        |  +-- Pseudunio_flabellata **      
        +--+                             
           +-- Pseudunio_flabellatiformis **
           |                             
           +-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
3313138822
  +----------- Cumberlandia          
--+                                  
  |  +-------- Margaritifera         
  +--+                               
     |  +----- Pseudunio_marocanus **   
     +--+                            
        |  +-- Pseudunio_auricularius **
        +--+                         
           +-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
3313197822
  +-------- Cumberlandia          
  |                               
--+-------- Margaritifera         
  |                               
  |  +----- Pseudunio_marocanus **   
  +--+                            
     |  +-- Pseudunio_auricularius **
     +--+                         
        +-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
3313628822
  +----- Pseudunio_marocanus **   
--+                            
  |  +-- Pseudunio_auricularius **
  +--+                         
     +-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
3313195822
  +----------- Margaritifera         
--+                                  
  |  +-------- Cumberlandia          
  +--+                               
     |  +----- Pseudunio_marocanus **   
     +--+                            
        |  +-- Pseudunio_auricularius **
        +--+                         
           +-- Pseudunio_homsensis **

Lopes-Lima et al., 2018 [Publ 3829], fig. 1

iTtTbiriSoTroTroSiD
331325822
  +----------- Margaritifera         
--+                                  
  |  +-------- Cumberlandia          
  +--+                               
     |  +----- Pseudunio_marocanus **   
     +--+                            
        |  +-- Pseudunio_auricularius **
        +--+                         
           +-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
3313197822
  +----------- Cumberlandia          
--+                                  
  |  +-------- Margaritifera         
  +--+                               
     |  +----- Pseudunio_marocanus **   
     +--+                            
        |  +-- Pseudunio_auricularius **
        +--+                         
           +-- Pseudunio_homsensis **

Lopes-Lima et al., 2018 [Publ 3829], fig. 3

iTtTbiriSoTroTroSiD
331325822
  +----------- Margaritifera         
--+                                  
  |  +-------- Cumberlandia          
  +--+                               
     |  +----- Pseudunio_marocanus **   
     +--+                            
        |  +-- Pseudunio_auricularius **
        +--+                         
           +-- Pseudunio_homsensis **

Chen et al., 2025 [Publ 4455], fig. 1

iTtTbiriSoTroTroSiD
315134238215
  +----------- Pseudunio_marocanus **      
  |                                     
  |        +-- Pseudunio_auricularius__1 **
--+        |                            
  |  +-----+-- Pseudunio_auricularius__2 **
  |  |     |                            
  +--+     +-- Pseudunio_homsensis **      
     |                                  
     |  +----- Gibbosula                
     +--+                               
        |  +-- Cumberlandia             
        +--+                            
           +-- Margaritifera

Vikhrev et al., 2018 [Publ 3812], fig. 3

iTtTbiriSoTroTroSiD
331310722
  +----------- Margaritifera         
--+                                  
  |  +-------- Cumberlandia          
  +--+                               
     |  +----- Pseudunio_marocanus **   
     +--+                            
        |  +-- Pseudunio_auricularius **
        +--+                         
           +-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
22027472--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Sano et al., 2020 [Publ 4138], fig. 3

iTtTbiriSoTroTroSiD
22026682--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Wu et al., 2020 [Publ 4121], fig. 2

iTtTbiriSoTroTroSiD
22023282--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
220213672--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Wu et al., 2020 [Publ 4121], fig. S1

iTtTbiriSoTroTroSiD
22023282--
  +-------- Margaritifera         
--+                               
  |  +----- Cumberlandia          
  +--+                            
     |  +-- Pseudunio_auricularius **
     +--+                         
        +-- Pseudunio_marocanus **

Bolotov et al., 2015 [Publ 3646], fig. 2B

iTtTbiriSoTroTroSiD
2202762--
  +-------- Cumberlandia          
--+                               
  |  +----- Margaritifera         
  +--+                            
     |  +-- Pseudunio_auricularius **
     +--+                         
        +-- Pseudunio_marocanus **

Sano et al., 2020 [Publ 4138], fig. 2

iTtTbiriSoTroTroSiD
22026682--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Bolotov et al., 2016 [Publ 3743], fig. 2

iTtTbiriSoTroTroSiD
22022872--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Inoue et al., 2013 [Publ 3579], fig. 3

iTtTbiriSoTroTroSiD
2202762--
  +-------- Cumberlandia          
--+                               
  |  +----- Margaritifera         
  +--+                            
     |  +-- Pseudunio_auricularius **
     +--+                         
        +-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
22023942--
  +-------- Cumberlandia          
--+                               
  |  +----- Margaritifera         
  +--+                            
     |  +-- Pseudunio_marocanus **   
     +--+                         
        +-- Pseudunio_auricularius **

Araujo et al., 2016 [Publ 3746], fig. 1_2b

iTtTbiriSoTroTroSiD
22021182--
  +-------- Margaritifera         
--+                               
  |  +----- Cumberlandia          
  +--+                            
     |  +-- Pseudunio_marocanus **   
     +--+                         
        +-- Pseudunio_auricularius **

Araujo et al., 2009 [Publ 2395], fig. 3

iTtTbiriSoTroTroSiD
2202652--
  +-------- Cumberlandia          
--+                               
  |  +----- Margaritifera         
  +--+                            
     |  +-- Pseudunio_marocanus **   
     +--+                         
        +-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
22027472--
  +----- Margaritifera         
--+                            
  |  +-- Pseudunio_auricularius **
  +--+                         
     +-- Pseudunio_marocanus **

Huang et al., 2018 [Publ 3828], fig. 3_5

iTtTbiriSoTroTroSiD
22022782--
  +-------- Margaritifera         
--+                               
  |  +----- Cumberlandia          
  +--+                            
     |  +-- Pseudunio_auricularius **
     +--+                         
        +-- Pseudunio_marocanus **

Sano et al., 2020 [Publ 4138], fig. 4

iTtTbiriSoTroTroSiD
22026682--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Takeuchi et al., 2015 [Publ 3661], fig. 2

iTtTbiriSoTroTroSiD
28028629
  +-------- Pseudunio_marocanus **   
--+                               
  |  +----- Pseudunio_auricularius **
  +--+                            
     |  +-- Cumberlandia          
     +--+                         
        +-- Margaritifera

Bolotov et al., 2016 [Publ 3743], fig. 4

iTtTbiriSoTroTroSiD
22022872--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Riccardi et al., 2020 [Publ 4089], fig. 2

iTtTbiriSoTroTroSiD
22021700--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
22023942--
  +-------- Cumberlandia          
--+                               
  |  +----- Margaritifera         
  +--+                            
     |  +-- Pseudunio_marocanus **   
     +--+                         
        +-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
220213672--
  +-- Pseudunio_auricularius **
--+                         
  +-- Pseudunio_marocanus **

Araujo et al., 2016 [Publ 3746], fig. 2a

iTtTbiriSoTroTroSiD
2202982--
  +-- Pseudunio_marocanus **   
--+                         
  +-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11012200--
-- Pseudunio_marocanus **

Zieritz et al., 2021 [Publ 4218], fig. 5

iTtTbiriSoTroTroSiD
11013011--
  +-- Margaritifera         
--+                         
  +-- Pseudunio_auricularius **

Bolotov et al., 2015 [Publ 3646], fig. 2A

iTtTbiriSoTroTroSiD
1101872--
  +----------- Margaritifera_margaritifera
  |                                       
  |     +----- Margaritifera_dahurica__1  
  |     |                                 
  +-----+----- Margaritifera_dahurica__2  
--+     |                                 
  |     |  +-- Pseudunio_auricularius **     
  |     +--+                              
  |        +-- Cumberlandia               
  |                                       
  |  +-------- Margaritifera_falcata      
  +--+                                    
     |  +----- Margaritifera_laevis       
     +--+                                 
        |  +-- Margaritifera_middendorffi 
        +--+                              
           +-- Margaritifera_marrianae

Bashê, 2022 [Publ 4265], fig. 1

iTtTbiriSoTroTroSiD
1101500--
-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
11014321--
  +-- Pseudunio_marocanus **
--+                      
  +-- Margaritifera

Walker et al., 2006 [Publ 2666], fig. 6

iTtTbiriSoTroTroSiD
11013552--
  +----- Cumberlandia          
--+                            
  |  +-- Pseudunio_auricularius **
  +--+                         
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11019542--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

Huff et al., 2004 [Publ 2697], fig. 4

iTtTbiriSoTroTroSiD
11011062--
  +-- Pseudunio_auricularius **
  |                         
--+-- Cumberlandia          
  |                         
  +-- Margaritifera

Huff et al., 2004 [Publ 2697], fig. 5-6

iTtTbiriSoTroTroSiD
11011162--
  +----- Margaritifera         
--+                            
  |  +-- Pseudunio_auricularius **
  +--+                         
     +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
11012122--
  +----- Cumberlandia       
--+                         
  |  +-- Margaritifera      
  +--+                      
     +-- Pseudunio_marocanus **

Araujo et al., 2009 [Publ 2589], fig. 7C

iTtTbiriSoTroTroSiD
1101900--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11017342--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

Gomes-dos-Santos et al., 2019 [Publ 3943], fig. 1-M

iTtTbiriSoTroTroSiD
1101522--
  +----- Margaritifera      
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Cumberlandia

Prie et al., 2012 [Publ 3487], fig. 2

iTtTbiriSoTroTroSiD
1101500--
-- Pseudunio_auricularius **

Araujo et al., 2009 [Publ 2589], fig. 7B

iTtTbiriSoTroTroSiD
11011000--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11017342--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11014321--
  +-- Pseudunio_marocanus **
--+                      
  +-- Margaritifera

Machordom et al., 2003 [Publ 2688], fig. 3

iTtTbiriSoTroTroSiD
1101322--
  +----- Cumberlandia          
--+                            
  |  +-- Margaritifera         
  +--+                         
     +-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11015542--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11011911--
  +-- Pseudunio_marocanus **
--+                      
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
11015311--
  +-- Pseudunio_marocanus **
--+                      
  +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
11019542--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

Zieritz et al., 2024 [Publ 4381], fig. 2

iTtTbiriSoTroTroSiD
11014411--
  +-- Margaritifera         
--+                         
  +-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11019052--
  +----- Margaritifera      
--+                         
  |  +-- Cumberlandia       
  +--+                      
     +-- Pseudunio_homsensis **

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

iTtTbiriSoTroTroSiD
11013721--
  +-- Pseudunio_marocanus **
--+                      
  +-- Margaritifera

Machordom et al., 2003 [Publ 2688], fig. 2

iTtTbiriSoTroTroSiD
1101211--
  +-- Margaritifera         
--+                         
  +-- Pseudunio_auricularius **

Araujo et al., 2017 [Publ 3801], fig. 2

iTtTbiriSoTroTroSiD
11011300--
-- Pseudunio_marocanus **

Araujo et al., 2009 [Publ 2589], fig. 7A

iTtTbiriSoTroTroSiD
11011000--
-- Pseudunio_auricularius **

Huff et al., 2004 [Publ 2697], fig. 3

iTtTbiriSoTroTroSiD
1101752--
  +-------- Margaritifera_margaritifera  
  |                                      
  |  +----- Cumberlandia                 
--+  |                                   
  |  |  +-- Pseudunio_auricularius **       
  |  |  |                                
  +--+  +-- Margaritifera_dahurica       
     |  |                                
     |  +-- Margaritifera_middendorffi__1
     |  |                                
     |  +-- Margaritifera_middendorffi__2
     +--+                                
        +-- Margaritifera_falcata__1     
        |                                
        +-- Margaritifera_falcata__2     
        |                                
        +-- Margaritifera_falcata__3     
        |                                
        +-- Margaritifera_falcata__4

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

iTtTbiriSoTroTroSiD
11012122--
  +----- Pseudunio_marocanus **
--+                         
  |  +-- Margaritifera      
  +--+                      
     +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
11014142--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11017342--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11013121--
  +-- Pseudunio_marocanus **
--+                      
  +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11014622--
  +----- Pseudunio_marocanus **
--+                         
  |  +-- Cumberlandia       
  +--+                      
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11011800--
-- Pseudunio_marocanus **

Gomes-dos-Santos et al., 2019 [Publ 3943], fig. 1

iTtTbiriSoTroTroSiD
1101942--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

Välilä et al., 2017 [Publ 3793], fig. 2

iTtTbiriSoTroTroSiD
1101211--
  +-- Margaritifera         
--+                         
  +-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11011800--
-- Pseudunio_marocanus **

Huff et al., 2004 [Publ 2697], fig. 2

iTtTbiriSoTroTroSiD
11011162--
  +-------- Margaritifera_dahurica        
  |                                       
  +-------- Margaritifera_margaritifera__1
  |                                       
  +-------- Margaritifera_margaritifera__2
  |                                       
--+-------- Margaritifera_margaritifera__3
  |                                       
  |     +-- Pseudunio_auricularius **        
  +-----+                                 
  |     +-- Cumberlandia                  
  |                                       
  |  +----- Margaritifera_falcata         
  +--+                                    
     |  +-- Margaritifera_middendorffi    
     +--+                                 
        +-- Margaritifera_marrianae

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

iTtTbiriSoTroTroSiD
11016842--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
1101900--
-- Pseudunio_marocanus **

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

iTtTbiriSoTroTroSiD
11018742--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

Takeuchi et al., 2015 [Publ 3661], fig. 3

iTtTbiriSoTroTroSiD
180297212
  +----- Pseudunio_auricularius__1 **
  |                               
--+----- Pseudunio_auricularius__2 **
  |                               
  |  +-- Cumberlandia             
  +--+                            
     +-- Margaritifera

Lopes-Lima et al., 2017 [Publ 3734], fig. 1

iTtTbiriSoTroTroSiD
11011611--
  +-- Pseudunio_auricularius **
--+                         
  +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11018542--
  +----- Cumberlandia       
--+                         
  |  +-- Pseudunio_marocanus **
  +--+                      
     +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11012211--
  +-- Pseudunio_marocanus **
--+                      
  +-- Margaritifera

Prié & Puillandre, 2014 [Publ 3529], fig. 2

iTtTbiriSoTroTroSiD
11011600--
-- Pseudunio_auricularius **

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

iTtTbiriSoTroTroSiD
11014721--
  +-- Pseudunio_marocanus **
--+                      
  +-- Margaritifera

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

iTtTbiriSoTroTroSiD
11019052--
  +----- Margaritifera      
--+                         
  |  +-- Pseudunio_homsensis **
  +--+                      
     +-- Cumberlandia

Araujo et al., 2018 [Publ 3779], fig. 1

iTtTbiriSoTroTroSiD
11011700--
-- Pseudunio_auricularius **
 
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