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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 | Unioninae | Unionini

subtribe UNIONINA Rafinesque, 1820: cladomics
     1 genus

Cladomics Summary

number of species = 226 (277 total)
number of subtaxa (rank: genus) = 1
analyzed in 231 cladograms in 136 publications
maximum number of terminals in an analysis = 26 (mean = 3.51, median = 3)
maximum number of subtaxa (genus) = 1 (mean = 1, median = 1)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 162
maximum number of included terminals = 26, (mean = 4.57 median = 3)
maximum number of subtaxa (genus) = 1 (mean = 1, median = 1)
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

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

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

iTtTbiriSoTroTroSiD
26262416064040
  +----------- Globunionina      
--+                              
  |  +-------- Aculamprotulina   
  +--+                           
     |  +----- Middendorffinaiina
     +--+                        
        |  +-- Unio **              
        +--+                     
           +-- Oxynaiina

Lyubas et al., 2022 [Publ 4255], fig. 7

iTtTbiriSoTroTroSiD
232320180028
1

Lyubas et al., 2022 [Publ 4255], fig. S1

iTtTbiriSoTroTroSiD
232314180022
1

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

iTtTbiriSoTroTroSiD
191917112127423
  +----------- Aculamprotulina   
--+                              
  |  +-------- Globunionina      
  +--+                           
     |  +----- Oxynaiina         
     +--+                        
        |  +-- Middendorffinaiina
        +--+                     
           +-- Unio **

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

iTtTbiriSoTroTroSiD
19191715629323
  +-------- Aculamprotulina   
--+                           
  |  +----- Unio **              
  +--+                        
     |  +-- Middendorffinaiina
     +--+                     
        +-- Oxynaiina

Lopes-Lima et al., 2024 [Publ 4376], fig. 4-5_MtG

iTtTbiriSoTroTroSiD
171715152121
  +-- Oxynaiina
--+            
  +-- Unio **

Lopes-Lima et al., 2024 [Publ 4376], fig. 4-5_AHE

iTtTbiriSoTroTroSiD
171715152131
  +-- Oxynaiina
--+            
  +-- Unio **

Lopes-Lima et al., 2021 [Publ 4217], fig. 1

iTtTbiriSoTroTroSiD
161614190025
1

Lopes-Lima et al., 2024 [Publ 4376], fig. 2

iTtTbiriSoTroTroSiD
141412100015
1

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

iTtTbiriSoTroTroSiD
13139151120
  +-- Oxynaiina
--+            
  +-- Unio **

Froufe et al., 2016 [Publ 3738], fig. 3

iTtTbiriSoTroTroSiD
121210122121
  +-- Oxynaiina
--+            
  +-- Unio **

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

iTtTbiriSoTroTroSiD
11119160024
1

Prie et al., 2026 [Publ 4818], fig. 2

iTtTbiriSoTroTroSiD
11119100021
1

Lopes-Lima et al., 2021 [Publ 4217], fig. 2

iTtTbiriSoTroTroSiD
10108100025
1

Khalloufi et al., 2011 [Publ 2748], fig. 2

iTtTbiriSoTroTroSiD
996120024
1

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

iTtTbiriSoTroTroSiD
995180026
1

Marrone et al., 2019 [Publ 3907], fig. 1

iTtTbiriSoTroTroSiD
884100023
1

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

iTtTbiriSoTroTroSiD
776140025
1

Reis et al., 2013 [Publ 3580], fig. 5

iTtTbiriSoTroTroSiD
776180024
1

Lopes-Lima et al., 2021 [Publ 4217], fig. 6

iTtTbiriSoTroTroSiD
775100022
1

Wu et al., 2024 [Publ 4379], fig. 5

iTtTbiriSoTroTroSiD
66414723223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
66414626223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
673151126
  +----------- Oxynaiina        
  |                             
  |        +-- Unio_gibbus **      
--+        |                    
  |  +-----+-- Unio_tumidus **     
  |  |     |                    
  |  |     +-- Anodontina       
  +--+                          
     |  +----- Unio_pictorum__1 ** 
     |  |                       
     |  +----- Unio_pictorum__2 ** 
     +--+                       
        +----- Unio_mancus **      
        |                       
        |  +-- Unio_tumidiformis **
        +--+                    
           +-- Unio_crassus **

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

iTtTbiriSoTroTroSiD
664180021
1

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

iTtTbiriSoTroTroSiD
66418516225
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
66418516225
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
66418916225
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
66318916224
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
664140024
1

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

iTtTbiriSoTroTroSiD
66413515223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
66418121323
  +-------- Globunionina   
--+                        
  |  +----- Aculamprotulina
  +--+                     
     |  +-- Unio **           
     +--+                  
        +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
553110024
1

Bashê, 2022 [Publ 4265], fig. 1

iTtTbiriSoTroTroSiD
554110027
1

Pfarrer et al., 2022 [Publ 4305], fig. 2-5

iTtTbiriSoTroTroSiD
553170024
1

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

iTtTbiriSoTroTroSiD
44216911223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Lopes-Lima et al., 2020 [Publ 4080], fig. 2-4

iTtTbiriSoTroTroSiD
441111824324
  +-------- Aculamprotulina   
--+                           
  |  +----- Middendorffinaiina
  +--+                        
     |  +-- Unio **              
     +--+                     
        +-- Oxynaiina

Wang et al., 2025 [Publ 4488], fig. 3B

iTtTbiriSoTroTroSiD
44216028323
  +-------- Aculamprotulina   
--+                           
  |  +----- Unio **              
  +--+                        
     |  +-- Middendorffinaiina
     +--+                     
        +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4421274223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

Wu et al., 2026 [Publ 4811], fig. 4

iTtTbiriSoTroTroSiD
44219128423
  +----------- Globunionina      
--+                              
  |  +-------- Aculamprotulina   
  +--+                           
     |  +----- Unio **              
     +--+                        
        |  +-- Middendorffinaiina
        +--+                     
           +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4421406223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

Klishko et al., 2018 [Publ 3945], fig. 8

iTtTbiriSoTroTroSiD
441144124
  +-- Oxynaiina
--+            
  +-- Unio **

Klishko et al., 2019 [Publ 3886], fig. 9

iTtTbiriSoTroTroSiD
441174224
  +----- Unio **              
--+                        
  |  +-- Middendorffinaiina
  +--+                     
     +-- Oxynaiina

Soroka, 2010 [Publ 3404], fig. 3b

iTtTbiriSoTroTroSiD
442150025
1

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

iTtTbiriSoTroTroSiD
44218123223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
44216911223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4421446223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
4421429223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4411777224
  +----- Unio **           
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
44216916223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Soroka, 2010 [Publ 3404], fig. 3a

iTtTbiriSoTroTroSiD
442150025
1

Wu et al., 2020 [Publ 4121], fig. 3

iTtTbiriSoTroTroSiD
4421449223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4421527223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
44214310223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Klymus et al., 2025 [Publ 4814], fig. 3B

iTtTbiriSoTroTroSiD
4421326223
1

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

iTtTbiriSoTroTroSiD
4421406223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

Nagel et al., 1998 [Publ 1946], fig. 8

iTtTbiriSoTroTroSiD
441100024
1

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

iTtTbiriSoTroTroSiD
442119516223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Nagel et al., 1998 [Publ 1946], fig. 1

iTtTbiriSoTroTroSiD
442150025
1

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

iTtTbiriSoTroTroSiD
442119717223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
44216411223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Soroka, 2005 [Publ 2726], fig. 3

iTtTbiriSoTroTroSiD
442110023
1

Liu et al., 2017 [Publ 3939], fig. 2

iTtTbiriSoTroTroSiD
4411117124
  +-- Unio **     
--+            
  +-- Oxynaiina

Wu et al., 2026 [Publ 4825], fig. 6B

iTtTbiriSoTroTroSiD
44216125323
  +-------- Aculamprotulina   
--+                           
  |  +----- Unio **              
  +--+                        
     |  +-- Middendorffinaiina
     +--+                     
        +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4421346223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

Soroka & Burzynski, 2017 [Publ 3837], fig. 1

iTtTbiriSoTroTroSiD
4421196223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
4421385223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
4421367223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
442161123
  +-- Aculamprotulina
--+                  
  +-- Unio **

Klishko et al., 2017 [Publ 3946], fig. 8

iTtTbiriSoTroTroSiD
442122123
  +-- Oxynaiina
--+            
  +-- Unio **

Kim et al., 2021 [Publ 4197], fig. 1

iTtTbiriSoTroTroSiD
4421148223
  +-- Unio **     
--+            
  +-- Oxynaiina

Wang et al., 2018 [Publ 3865], fig. 2

iTtTbiriSoTroTroSiD
4421273223
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

Wu et al., 2022 [Publ 4238], fig. 2d

iTtTbiriSoTroTroSiD
44213419223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4421161123
  +-- Aculamprotulina
--+                  
  +-- Unio **

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

iTtTbiriSoTroTroSiD
442119717223
1

Klymus et al., 2025 [Publ 4814], fig. 3A

iTtTbiriSoTroTroSiD
4421326223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Fukata & Iigo, 2019 [Publ 4108], fig. 1

iTtTbiriSoTroTroSiD
4421139223
  +-- Unio **     
--+            
  +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
4421509223
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Bogatov et al., 2018 [Publ 3859], fig. 1

iTtTbiriSoTroTroSiD
353130026
  +-------- Unio_tumidus ** 
--+                      
  |  +----- Anodontina   
  +--+                   
     |  +-- Unio_crassus ** 
     +--+                
        +-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
3311160024
1

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

iTtTbiriSoTroTroSiD
3311382224
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
3311737224
  +----- Unio **           
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
3311737224
  +----- Unio **           
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Oxynaiina

Soroka, 2010 [Publ 3404], fig. 3b-M

iTtTbiriSoTroTroSiD
331130024
1

Wu et al., 2022 [Publ 4238], fig. 2e

iTtTbiriSoTroTroSiD
33111510224
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
3311202224
  +----- Oxynaiina      
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
3311150024
1

Bogatov et al., 2018 [Publ 3859], fig. 2

iTtTbiriSoTroTroSiD
353130028
  +-------- Unio_crassus ** 
--+                      
  |  +----- Unio_pictorum **
  +--+                   
     |  +-- Unio_tumidus ** 
     +--+                
        +-- Anodontina

Froufe et al., 2017 [Publ 3802], fig. 7

iTtTbiriSoTroTroSiD
331170024
1

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

iTtTbiriSoTroTroSiD
33116610224
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Källersjö et al., 2005 [Publ 2684], fig. 1A

iTtTbiriSoTroTroSiD
330140025
1

Dyduch-Falniowska & Koziol, 1989 [Publ 3814], fig. 5_7

iTtTbiriSoTroTroSiD
331120026
1

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

iTtTbiriSoTroTroSiD
3311160024
1

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

iTtTbiriSoTroTroSiD
3311382224
  +-- Oxynaiina
--+            
  +-- Unio **

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

iTtTbiriSoTroTroSiD
33116610224
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
33111367224
  +----- Unio **           
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Oxynaiina

Soroka, 2010 [Publ 3404], fig. 2

iTtTbiriSoTroTroSiD
330140025
1

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

iTtTbiriSoTroTroSiD
33116610224
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
3311792124
  +-- Unio **     
--+            
  +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
3311192224
  +----- Aculamprotulina
--+                     
  |  +-- Oxynaiina      
  +--+                  
     +-- Unio **

Konopleva et al., 2023 [Publ 4269], fig. S1_4

iTtTbiriSoTroTroSiD
3311539324
  +-------- Middendorffinaiina
--+                           
  |  +----- Aculamprotulina   
  +--+                        
     |  +-- Unio **              
     +--+                     
        +-- Oxynaiina

Soroka, 2010 [Publ 3404], fig. 2-M

iTtTbiriSoTroTroSiD
331120026
1

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

iTtTbiriSoTroTroSiD
3311192224
  +----- Oxynaiina      
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Unio **

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

iTtTbiriSoTroTroSiD
310011227232
  +-- Unio_crassus **         
  |                        
  +-- Unio_pictorum **        
  |                        
  +-- Unio_tumidus **         
  |                        
  +-- Aculamprotulina      
  |                        
  +-- Cuneopsis_szechenyii 
--+                        
  +-- Tchangsinaia         
  |                        
  +-- Cuneopsis_rufescens  
  |                        
  +-- Nodularia_douglasiae 
  |                        
  +-- Nodularia_nuxpersicae
  |                        
  +-- Schistodesmus

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

iTtTbiriSoTroTroSiD
3311141124
  +-- Aculamprotulina
--+                  
  +-- Unio **

Soroka, 2010 [Publ 3404], fig. 4

iTtTbiriSoTroTroSiD
331140024
1

Manendo et al., 2008 [Publ 2669], fig. 1

iTtTbiriSoTroTroSiD
3311100026
1

Soroka, 2010 [Publ 3404], fig. 8a

iTtTbiriSoTroTroSiD
331130024
1

Källersjö et al., 2005 [Publ 2684], fig. 1B-F

iTtTbiriSoTroTroSiD
331140024
1

Soroka, 2010 [Publ 3404], fig. 8b

iTtTbiriSoTroTroSiD
331130024
1

Zieritz et al., 2012 [Publ 3673], fig. 4

iTtTbiriSoTroTroSiD
331150026
1

Kilikowska et al., 2020 [Publ 4335], fig. 4a

iTtTbiriSoTroTroSiD
331100024
1

Soroka, 2010 [Publ 3404], fig. 5b

iTtTbiriSoTroTroSiD
331150024
1

Soroka, 2010 [Publ 3404], fig. 3a-M

iTtTbiriSoTroTroSiD
331130026
1

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

iTtTbiriSoTroTroSiD
33111367224
  +----------- Unio **                
  |                                
--+  +-------- Aculamprotulina     
  |  |                             
  |  +-------- Schistodesmus       
  +--+                             
     +-------- Nodularia           
     |                             
     |  +----- Cuneopsis_szechenyii
     +--+                          
        |  +-- Tchangsinaia        
        +--+                       
           +-- Cuneopsis_rufescens

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

iTtTbiriSoTroTroSiD
3311332224
  +----- Middendorffinaiina
--+                        
  |  +-- Oxynaiina         
  +--+                     
     +-- Unio **

Soroka, 2010 [Publ 3404], fig. 5a

iTtTbiriSoTroTroSiD
331150026
1

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

iTtTbiriSoTroTroSiD
3311200024
1

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

iTtTbiriSoTroTroSiD
2501290028
  +-------- Unio_pictorum **
--+                      
  |  +----- Unio_tumidus ** 
  +--+                   
     |  +-- Anodontina   
     +--+                
        +-- Alasmidontina

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

iTtTbiriSoTroTroSiD
220142194--
  +-------------------- Globunionina      
  |                                       
--+  +----------------- Aculamprotulina   
  |  |                                    
  |  |              +-- Middendorffinaiina
  +--+  +-----------+                     
     |  |           +-- Nodularia         
     |  |                                 
     +--+  +----------- Cuneopsis         
        |  |                              
        |  |        +-- Diaurora          
        +--+--------+                     
           |        +-- Schistodesmus     
           |                              
           |  +-------- Pseudocuneopsis   
           +--+                           
              |  +----- Inversiunio       
              +--+                        
                 |  +-- Tchangsinaia      
                 +--+                     
                    +-- Unio **

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

iTtTbiriSoTroTroSiD
22014500--
1

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

iTtTbiriSoTroTroSiD
22014400--
1

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

iTtTbiriSoTroTroSiD
22011700--
1

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

iTtTbiriSoTroTroSiD
220139163--
  +----------------- Aculamprotulina   
  |                                    
  |              +-- Middendorffinaiina
--+  +-----------+                     
  |  |           +-- Nodularia         
  |  |                                 
  +--+  +----------- Cuneopsis         
     |  |                              
     |  |        +-- Diaurora          
     +--+--------+                     
        |        +-- Schistodesmus     
        |                              
        |  +-------- Pseudocuneopsis   
        +--+                           
           |  +----- Inversiunio       
           +--+                        
              |  +-- Tchangsinaia      
              +--+                     
                 +-- Unio **

Tomilova et al., 2020 [Publ 4097], fig. 3

iTtTbiriSoTroTroSiD
2201600--
1

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

iTtTbiriSoTroTroSiD
220151122--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
2201900--
1

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

iTtTbiriSoTroTroSiD
220139163--
  +----------------- Aculamprotulina   
  |                                    
  |              +-- Middendorffinaiina
--+  +-----------+                     
  |  |           +-- Nodularia         
  |  |                                 
  +--+  +----------- Cuneopsis         
     |  |                              
     |  |        +-- Diaurora          
     +--+--------+                     
        |        +-- Schistodesmus     
        |                              
        |  +-------- Pseudocuneopsis   
        +--+                           
           |  +----- Inversiunio       
           +--+                        
              |  +-- Tchangsinaia      
              +--+                     
                 +-- Unio **

Sonowal et al., 2021 [Publ 4221], fig. 3b

iTtTbiriSoTroTroSiD
2901131132
  +-- Aculamprotulina
  |                  
  +-- Lamprotula     
  |                  
  +-- Lampsilis      
  |                  
  +-- Potomida       
  |                  
--+-- Unio_pictorum **  
  |                  
  +-- Anodontina     
  |                  
  +-- Unio_ravoisieri **
  |                  
  +-- Lamellidens    
  |                  
  +-- Elliptio

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
22011600--
1

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

iTtTbiriSoTroTroSiD
22012900--
1

Soroka, 2010 [Publ 3404], fig. 8a-M

iTtTbiriSoTroTroSiD
2201100--
1

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

iTtTbiriSoTroTroSiD
22013400--
1

Chen et al., 2025 [Publ 4487], fig. 3

iTtTbiriSoTroTroSiD
220139163--
  +-------- Globunionina   
--+                        
  |  +----- Aculamprotulina
  +--+                     
     |  +-- Unio **           
     +--+                  
        +-- Oxynaiina

Sonowal et al., 2021 [Publ 4221], fig. 3a

iTtTbiriSoTroTroSiD
2401131127
  +-- Anodontina     
  |                  
  +-- Unio_ravoisieri **
--+                  
  +-- Unio_pictorum **  
  |                  
  +-- Aculamprotulina

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

iTtTbiriSoTroTroSiD
22011111--
  +-- Aculamprotulina
--+                  
  +-- Unio **

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

iTtTbiriSoTroTroSiD
22012800--
1

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

iTtTbiriSoTroTroSiD
220139163--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
22012800--
1

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

iTtTbiriSoTroTroSiD
22012800--
1

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

iTtTbiriSoTroTroSiD
2701231131
  +-------- Unio_pictorum__2 **
  |                         
--+     +-- Unio_pictorum__1 **
  |  +--+                   
  |  |  +-- Unio_caffer **     
  +--+                      
     |  +-- Anodontina      
     +--+                   
        +-- Alasmidontina

Wu et al., 2025 [Publ 4442], fig. 6

iTtTbiriSoTroTroSiD
22011500--
1

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

iTtTbiriSoTroTroSiD
22013700--
1

Sayenko et al., 2021 [Publ 4189], fig. 6

iTtTbiriSoTroTroSiD
220111112--
  +-------- Unio **              
--+                           
  |  +----- Inversiunio       
  +--+                        
     |  +-- Middendorffinaiina
     +--+                     
        +-- Nodularia

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

iTtTbiriSoTroTroSiD
22012952--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Bogan & Hoeh, 2000 [Publ 2650], fig. 1

iTtTbiriSoTroTroSiD
22013200--
1

Soroka, 2010 [Publ 3404], fig. 8b-M

iTtTbiriSoTroTroSiD
2201100--
1

Sonowal et al., 2021 [Publ 4221], fig. 4

iTtTbiriSoTroTroSiD
2401141127
  +----------- Aculamprotulina
--+                           
  |  +-------- Unio_pictorum **  
  +--+                        
     |  +----- Anodontina     
     +--+                     
        |  +-- Unio_ravoisieri **
        +--+                  
           +-- Lamellidens

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

iTtTbiriSoTroTroSiD
22012800--
1

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

iTtTbiriSoTroTroSiD
2501290028
  +----- Unio_pictorum **
  |                   
--+----- Unio_tumidus ** 
  |                   
  |  +-- Anodontina   
  +--+                
     +-- Alasmidontina

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

iTtTbiriSoTroTroSiD
22018992--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
22018992--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
22013052--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
22012900--
1

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

iTtTbiriSoTroTroSiD
22012900--
1

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

iTtTbiriSoTroTroSiD
22011111--
  +-- Aculamprotulina
--+                  
  +-- Unio **

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

iTtTbiriSoTroTroSiD
220142194--
  +----------------------- Globunionina      
  |                                          
--+  +-------------------- Aculamprotulina   
  |  |                                       
  +--+  +----------------- Middendorffinaiina
     |  |                                    
     +--+  +-------------- Nodularia         
        |  |                                 
        +--+  +----------- Cuneopsis         
           |  |                              
           |  |        +-- Diaurora          
           +--+--------+                     
              |        +-- Schistodesmus     
              |                              
              |  +-------- Pseudocuneopsis   
              +--+                           
                 |  +----- Unio **              
                 +--+                        
                    |  +-- Tchangsinaia      
                    +--+                     
                       +-- Inversiunio

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

iTtTbiriSoTroTroSiD
2201900--
1

Hoeh et al., 2009 [Publ 2716], fig. 3

iTtTbiriSoTroTroSiD
22014500--
1

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

iTtTbiriSoTroTroSiD
220151162--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Hoeh et al., 2009 [Publ 2716], fig. 4

iTtTbiriSoTroTroSiD
22014400--
1

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

iTtTbiriSoTroTroSiD
22013400--
1

Chase et al., 2018 [Publ 3847], fig. 5

iTtTbiriSoTroTroSiD
2201800--
1

Bieler et al., 2014 [Publ 3622], fig. 26_27_29_30

iTtTbiriSoTroTroSiD
1101400--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
1101800--
-- Unio_delphinus **

Graf et al., 2015 [Publ 3619], fig. 1

iTtTbiriSoTroTroSiD
11011900--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
1101900--
-- Unio_delphinus **

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

iTtTbiriSoTroTroSiD
11012411--
  +-- Aculamprotulina
--+                  
  +-- Unio **

Dai et al., 2024 [Publ 4386], fig. 5-BI

iTtTbiriSoTroTroSiD
110121113--
  +----- Unio **              
--+                        
  |  +-- Middendorffinaiina
  +--+                     
     +-- Oxynaiina

Combosch et al., 2017 [Publ 3806], fig. 1

iTtTbiriSoTroTroSiD
11011400--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11012100--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11011500--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11011100--
-- Unio_ravoisieri **

Combosch et al., 2017 [Publ 3806], fig. 2

iTtTbiriSoTroTroSiD
11011400--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11012700--
-- Unio_pictorum **

Graf et al., 2014 [Publ 3591], fig. 3

iTtTbiriSoTroTroSiD
11011000--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11015621--
  +-- Unio **     
--+            
  +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
11011521--
-- Unio_pictorum **

Graf, 2000 [Publ 2647], fig. 1

iTtTbiriSoTroTroSiD
11011600--
-- Unio **

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

iTtTbiriSoTroTroSiD
1101900--
-- Unio_delphinus **

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

iTtTbiriSoTroTroSiD
11012111--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11011200--
-- Unio_ravoisieri **

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

iTtTbiriSoTroTroSiD
1101611--
  +-- Unio **           
--+                  
  +-- Aculamprotulina

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

iTtTbiriSoTroTroSiD
11013341--
  +-- Unio **           
--+                  
  +-- Aculamprotulina

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

iTtTbiriSoTroTroSiD
11012042--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
11013662--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

Bieler et al., 2014 [Publ 3622], fig. 28

iTtTbiriSoTroTroSiD
1101400--
-- Unio_pictorum **

Nagel et al., 1998 [Publ 1946], fig. 9

iTtTbiriSoTroTroSiD
1101500--
-- Unio **

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

iTtTbiriSoTroTroSiD
11011800--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11013862--
  +-- Unio **     
--+            
  +-- Oxynaiina

González & Giribet, 2015 [Publ 3632], fig. 3

iTtTbiriSoTroTroSiD
1101500--
-- Unio_pictorum **

Chong et al., 2008 [Publ 2668], fig. 3

iTtTbiriSoTroTroSiD
11011700--
-- Unio_tumidus **

Taylor et al., 2007 [Publ 2696], fig. 1-3

iTtTbiriSoTroTroSiD
1101300--
-- Unio_pictorum **

Liu et al., 2023 [Publ 4310], fig. 4

iTtTbiriSoTroTroSiD
110123132--
  +----- Unio **           
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
11014400--
-- Unio_crassus **

González & Giribet, 2015 [Publ 3632], fig. 2

iTtTbiriSoTroTroSiD
1101500--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11013341--
  +-- Unio **           
--+                  
  +-- Aculamprotulina

Liu et al., 2023 [Publ 4310], fig. 5

iTtTbiriSoTroTroSiD
110123132--
  +-- Unio **           
  |                  
--+-- Aculamprotulina
  |                  
  +-- Oxynaiina

Graf & Ó Foighil, 2000 [Publ 2649], fig. 1_2

iTtTbiriSoTroTroSiD
11011000--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
1101500--
-- Unio_delphinus **

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

iTtTbiriSoTroTroSiD
1101800--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11011322--
-- Unio_delphinus **

Choi et al., 2020 [Publ 4169], fig. 2a

iTtTbiriSoTroTroSiD
110111111--
  +----------- Nodularia    
--+                         
  |  +-------- Schistodesmus
  +--+                      
     |  +----- Cuneopsis    
     +--+                   
        |  +-- Unio **         
        +--+                
           +-- Tchangsinaia

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

iTtTbiriSoTroTroSiD
11012411--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
1101611--
  +-- Unio **           
--+                  
  +-- Aculamprotulina

Marshall et al., 2014 [Publ 3593], fig. 1

iTtTbiriSoTroTroSiD
11011300--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11011432--
  +-------- Aculamprotulina
--+                        
  |  +----- Schistodesmus  
  +--+                     
     |  +-- Nodularia      
     +--+                  
        +-- Unio **

Dai et al., 2024 [Publ 4386], fig. 5-ML

iTtTbiriSoTroTroSiD
110121113--
  +-------- Aculamprotulina   
--+                           
  |  +----- Unio **              
  +--+                        
     |  +-- Middendorffinaiina
     +--+                     
        +-- Oxynaiina

Choi et al., 2020 [Publ 4169], fig. 3a

iTtTbiriSoTroTroSiD
1101551--
  +----- Nodularia
--+               
  |  +-- Cuneopsis
  +--+            
     +-- Unio **

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

iTtTbiriSoTroTroSiD
110129102--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

González & Giribet, 2015 [Publ 3632], fig. 4

iTtTbiriSoTroTroSiD
1101500--
-- Unio_pictorum **

Klunzinger et al., 2021 [Publ 4086], fig. 2

iTtTbiriSoTroTroSiD
1101500--
-- Unio_pictorum **

Wu et al., 2020 [Publ 4147], fig. 4

iTtTbiriSoTroTroSiD
11011341--
  +-- Unio **           
--+                  
  +-- Aculamprotulina

Wang et al., 2016 [Publ 3714], fig. 2

iTtTbiriSoTroTroSiD
1101200--
-- Unio_tumidus **

Giribet & Distel, 2003 [Publ 3670], fig. 3f5

iTtTbiriSoTroTroSiD
1101500--
-- Unio_ravoisieri **

González & Giribet, 2015 [Publ 3632], fig. 6

iTtTbiriSoTroTroSiD
1101500--
-- Unio_pictorum **

Wu et al., 2026 [Publ 4811], fig. 3

iTtTbiriSoTroTroSiD
11011631--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11015621--
  +-- Unio **     
--+            
  +-- Oxynaiina

Dai et al., 2024 [Publ 4386], fig. 3

iTtTbiriSoTroTroSiD
110113103--
-- Unio_crassus **

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

iTtTbiriSoTroTroSiD
11012152--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
11011100--
-- Unio_ravoisieri **

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

iTtTbiriSoTroTroSiD
11013042--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
1101800--
-- Unio_pictorum **

Wu et al., 2023 [Publ 4296], fig. 4

iTtTbiriSoTroTroSiD
110122122--
  +----- Unio **           
--+                     
  |  +-- Aculamprotulina
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
11011911--
  +-- Oxynaiina
--+            
  +-- Unio **

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

iTtTbiriSoTroTroSiD
11015262--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
11012811--
  +-- Unio **           
--+                  
  +-- Aculamprotulina

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

iTtTbiriSoTroTroSiD
11015282--
  +----- Aculamprotulina
--+                     
  |  +-- Unio **           
  +--+                  
     +-- Oxynaiina

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

iTtTbiriSoTroTroSiD
11012700--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
11013900--
-- Unio_pictorum **

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

iTtTbiriSoTroTroSiD
1101700--
-- Unio **

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

iTtTbiriSoTroTroSiD
11011000--
-- Unio_ravoisieri **
 
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