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Page last updated
Thu 17 July 2025

The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)

FM(U)otW(aolcb) is the web version of the MUSSEL Project Database. Follow the links to browse the data or use the custom Google search field. Either way, you win!


Bivalvia | Unionoida | Unionoidea | Unionidae | Gonideinae | Contradentini

genus Lens Simpson, 1900: cladomics
     13 Recent valid species (13 total species)

Cladomics Summary

number of species = 13 (13 total)
analyzed in 64 cladograms in 41 publications
maximum number of terminals in an analysis = 11 (mean = 2.84, median = 1.5)

Basic Monophyly

number of cladograms that test basic monophyly (i.e., more than 1 included species) = 32
maximum number of included terminals = 11, (mean = 4.69 median = 4)
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


Jeratthitikul et al., 2019 [Publ 3911], fig. 4

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
449175210
  +----------------------- Physunio           
  |                                           
  |                    +-- Lens_maenamensis **   
--+  +-----------------+                      
  |  |                 +-- Lens_rolfbrandti **   
  |  |                                        
  |  |  +----------------- Lens_contradens__12 **
  +--+  |                                     
     |  |           +----- Lens_micropterus__3 **
     |  |           |                         
     |  +-----------+  +-- Lens_contradens__10 **
     |  |           |  |                      
     +--+           +--+-- Lens_micropterus__2 **
        |              |                      
        |              +-- Lens_contradens__11 **
        |                                     
        |  +-------------- Lens_contradens__9 ** 
        |  |                                  
        +--+  +----------- Lens_contradens__8 ** 
           |  |                               
           +--+  +-------- Lens_contradens__7 ** 
              |  |                            
              +--+  +----- Lens_contradens__6 ** 
                 |  |                         
                 |  |  +-- Lens_contradens__1 ** 
                 +--+  |                      
                    |  +-- Lens_contradens__2 ** 
                    |  |                      
                    |  +-- Lens_contradens__3 ** 
                    +--+                      
                       +-- Lens_contradens__4 ** 
                       |                      
                       +-- Lens_micropterus__1 **
                       |                      
                       +-- Lens_contradens__5 ** 

Pfeiffer et al., 2021 [Publ 4126], fig. 9

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
8868217415
                    +-- Trapezoideus    
  +-----------------+                   
  |                 +-- Physunio        
  |                                     
--+  +----------------- Yaukthwa        
  |  |                                  
  |  |  +-------------- Solenaia        
  +--+  |                               
     |  |     +-------- Lens_misellus **   
     |  |  +--+                         
     +--+  |  |  +----- Lens_inornatus **  
        |  |  +--+                      
        |  |     |  +-- Lens_contradens ** 
        |  |     +--+                   
        +--+        +-- Lens_micropterus **
           |                            
           |        +-- Lens_maenamensis **
           |     +--+                   
           |     |  +-- Lens_eximius **    
           +-----+                      
                 |  +-- Lens_pallegoixi ** 
                 +--+                   
                    +-- Lens_comptus **    

Zieritz et al., 2024 [Publ 4381], fig. 7A

iTtTbiriSoTroTroSiD
111191111114
  +----------------- Solenaia        
  |                                  
  |           +----- Lens_contradens ** 
--+  +--------+                      
  |  |        |  +-- Lens_lugens **     
  |  |        +--+                   
  |  |           +-- Lens_micropterus **
  +--+                               
     |           +-- Lens_inornatus **  
     |  +--------+                   
     |  |        +-- Lens_misellus **   
     |  |                            
     +--+     +----- Lens_maenamensis **
        |  +--+                      
        |  |  |  +-- Lens_eximius **    
        |  |  +--+                   
        +--+     +-- Lens_rolfbrandti **
           |                         
           |  +----- Lens_pallegoixi ** 
           +--+                      
              |  +-- Lens_comptus **    
              +--+                   
                 +-- Lens_novoselovi ** 

Pfeiffer et al., 2021 [Publ 4126], fig. 12

iTtTbiriSoTroTroSiD
101071011114
  +-------------- Solenaia        
  |                               
  |           +-- Lens_contradens ** 
  |  +--------+                   
--+  |        +-- Lens_micropterus **
  |  |                            
  |  |        +-- Lens_inornatus **  
  |  +--------+                   
  +--+        +-- Lens_misellus **   
     |                            
     |     +----- Lens_maenamensis **
     |  +--+                      
     |  |  |  +-- Lens_rolfbrandti **
     |  |  +--+                   
     +--+     +-- Lens_eximius **    
        |                         
        |  +----- Lens_pallegoixi ** 
        +--+                      
           |  +-- Lens_comptus **    
           +--+                   
              +-- Lens_novoselovi ** 

Bolotov et al., 2020 [Publ 4100], fig. S1

iTtTbiriSoTroTroSiD
997910314314
  +-------------------- Physunio        
  |                                     
--+  +----------------- Trapezoideus    
  |  |                                  
  +--+  +-------------- Yaukthwa        
     |  |                               
     |  |        +----- Lens_misellus **   
     +--+  +-----+                      
        |  |     |  +-- Lens_contradens ** 
        |  |     +--+                   
        |  |        +-- Lens_micropterus **
        +--+                            
           |     +----- Lens_maenamensis **
           |  +--+                      
           |  |  |  +-- Lens_eximius **    
           |  |  +--+                   
           +--+     +-- Lens_rolfbrandti **
              |                         
              |  +----- Lens_pallegoixi ** 
              +--+                      
                 |  +-- Lens_novoselovi ** 
                 +--+                   
                    +-- Lens_comptus **    

Konopleva et al., 2019 [Publ 4041], fig. 3

iTtTbiriSoTroTroSiD
99792510314
  +----------------- Yaukthwa        
  |                                  
  |              +-- Trapezoideus    
--+  +-----------+                   
  |  |           +-- Physunio        
  |  |                               
  |  |        +----- Lens_misellus **   
  +--+  +-----+                      
     |  |     |  +-- Lens_contradens ** 
     |  |     +--+                   
     |  |        +-- Lens_micropterus **
     +--+                            
        |     +----- Lens_pallegoixi ** 
        |  +--+                      
        |  |  |  +-- Lens_novoselovi ** 
        |  |  +--+                   
        +--+     +-- Lens_comptus **    
           |                         
           |  +----- Lens_maenamensis **
           +--+                      
              |  +-- Lens_eximius **    
              +--+                   
                 +-- Lens_rolfbrandti **

Konopleva et al., 2023 [Publ 4262], fig. 8

iTtTbiriSoTroTroSiD
99797310314
                 +-- Physunio        
  +--------------+                   
  |              +-- Trapezoideus    
  |                                  
--+  +-------------- Yaukthwa        
  |  |                               
  |  |        +----- Lens_misellus **   
  +--+  +-----+                      
     |  |     |  +-- Lens_contradens ** 
     |  |     +--+                   
     |  |        +-- Lens_micropterus **
     +--+                            
        |     +----- Lens_pallegoixi ** 
        |  +--+                      
        |  |  |  +-- Lens_novoselovi ** 
        |  |  +--+                   
        +--+     +-- Lens_comptus **    
           |                         
           |  +----- Lens_maenamensis **
           +--+                      
              |  +-- Lens_eximius **    
              +--+                   
                 +-- Lens_rolfbrandti **

Bolotov et al., 2020 [Publ 4100], fig. 1

iTtTbiriSoTroTroSiD
997910914314
                 +-- Trapezoideus    
  +--------------+                   
  |              +-- Physunio        
  |                                  
--+  +-------------- Yaukthwa        
  |  |                               
  |  |        +----- Lens_misellus **   
  +--+  +-----+                      
     |  |     |  +-- Lens_contradens ** 
     |  |     +--+                   
     |  |        +-- Lens_micropterus **
     +--+                            
        |     +----- Lens_pallegoixi ** 
        |  +--+                      
        |  |  |  +-- Lens_comptus **    
        |  |  +--+                   
        +--+     +-- Lens_novoselovi ** 
           |                         
           |  +----- Lens_maenamensis **
           +--+                      
              |  +-- Lens_eximius **    
              +--+                   
                 +-- Lens_rolfbrandti **

Konopleva et al., 2019 [Publ 3900], fig. 1

iTtTbiriSoTroTroSiD
7757459316
  +----------------------- Yaukthwa        
  |                                        
--+  +-------------------- Physunio        
  |  |                                     
  +--+  +----------------- Trapezoideus    
     |  |                                  
     +--+              +-- Lens_contradens ** 
        |  +-----------+                   
        |  |           +-- Lens_micropterus **
        +--+                               
           |  +----------- Lens_misellus **   
           +--+                            
              |  +-------- Lens_eximius **    
              +--+                         
                 |  +----- Lens_pallegoixi ** 
                 +--+                      
                    |  +-- Lens_comptus **    
                    +--+                   
                       +-- Lens_novoselovi ** 

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

iTtTbiriSoTroTroSiD
612761936217
  +----------- Physunio_modelli 
  |                             
  +----------- Lens **             
  |                             
  +----------- Physunio_superbus
--+                             
  +----------- Yaukthwa         
  |                             
  |        +-- Lens_comptus **     
  |  +-----+                    
  |  |     +-- Lens_novoselovi **  
  +--+                          
     |  +----- Lens_eximius **     
     +--+                       
        |  +-- Lens_contradens **  
        +--+                    
           +-- Lens_micropterus ** 

Zieritz et al., 2020 [Publ 4102], fig. 4

iTtTbiriSoTroTroSiD
6646258318
  +-------------------- Physunio       
  |                                    
--+  +----------------- Yaukthwa       
  |  |                                 
  +--+  +-------------- Trapezoideus   
     |  |                              
     +--+           +-- Lens_contradens **
        |  +--------+                  
        |  |        +-- Lens_misellus **  
        +--+                           
           |  +-------- Lens_eximius **   
           +--+                        
              |  +----- Lens_pallegoixi **
              +--+                     
                 |  +-- Lens_comptus **   
                 +--+                  
                    +-- Lens_novoselovi **

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

iTtTbiriSoTroTroSiD
55351347216
  +-------------- Physunio        
  |                               
--+  +----------- Yaukthwa        
  |  |                            
  +--+        +-- Lens_contradens ** 
     |  +-----+                   
     |  |     +-- Lens_micropterus **
     +--+                         
        |  +----- Lens_eximius **    
        +--+                      
           |  +-- Lens_comptus **    
           +--+                   
              +-- Lens_novoselovi ** 

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

iTtTbiriSoTroTroSiD
55351947216
  +-------------- Physunio        
  |                               
--+  +----------- Yaukthwa        
  |  |                            
  +--+        +-- Lens_contradens ** 
     |  +-----+                   
     |  |     +-- Lens_micropterus **
     +--+                         
        |  +----- Lens_eximius **    
        +--+                      
           |  +-- Lens_comptus **    
           +--+                   
              +-- Lens_novoselovi ** 

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

iTtTbiriSoTroTroSiD
55351207216
  +-------------- Physunio        
  |                               
--+  +----------- Yaukthwa        
  |  |                            
  +--+        +-- Lens_contradens ** 
     |  +-----+                   
     |  |     +-- Lens_micropterus **
     +--+                         
        |  +----- Lens_eximius **    
        +--+                      
           |  +-- Lens_comptus **    
           +--+                   
              +-- Lens_novoselovi ** 

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

iTtTbiriSoTroTroSiD
55351927216
  +-------------- Physunio        
  |                               
--+  +----------- Yaukthwa        
  |  |                            
  +--+        +-- Lens_contradens ** 
     |  +-----+                   
     |  |     +-- Lens_micropterus **
     +--+                         
        |  +----- Lens_eximius **    
        +--+                      
           |  +-- Lens_novoselovi ** 
           +--+                   
              +-- Lens_comptus **    

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

iTtTbiriSoTroTroSiD
51005785216
  +-- Lens_eximius **       
  |                      
  +-- Physunio_modelli__1
  |                      
  +-- Yaukthwa_nesemanni 
  |                      
  +-- Yaukthwa_panhai    
  |                      
  +-- Lens_comptus **       
  |                      
--+-- Lens_novoselovi **    
  |                      
  +-- Physunio_modelli__2
  |                      
  +-- Physunio           
  |                      
  +-- Yaukthwa_inlenensis
  |                      
  +-- Lens_contradens **    
  |                      
  +-- Lens_micropterus **   

Jeratthitikul et al., 2021 [Publ 4212], fig. 4

iTtTbiriSoTroTroSiD
4424176315
              +-- Trapezoideus    
  +-----------+                   
  |           +-- Physunio        
--+                               
  |  +----------- Yaukthwa        
  +--+                            
     |  +-------- Lens_contradens ** 
     +--+                         
        |  +----- Lens_novoselovi ** 
        +--+                      
           |  +-- Lens_rolfbrandti **
           +--+                   
              +-- Lens_maenamensis **

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

iTtTbiriSoTroTroSiD
44241357217
           +-- Physunio       
  +--------+                  
  |        +-- Yaukthwa       
--+                           
  |  +-------- Lens_contradens **
  +--+                        
     |  +----- Lens_eximius **   
     +--+                     
        |  +-- Lens_comptus **   
        +--+                  
           +-- Lens_novoselovi **

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

iTtTbiriSoTroTroSiD
3313435317
           +-- Physunio       
  +--------+                  
  |        +-- Trapezoideus   
--+                           
  |  +-------- Yaukthwa       
  +--+                        
     |  +----- Lens_eximius **   
     +--+                     
        |  +-- Lens_contradens **
        +--+                  
           +-- Lens_lugens **    

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

iTtTbiriSoTroTroSiD
22023053--
  +----------- Physunio       
--+                           
  |  +-------- Trapezoideus   
  +--+                        
     |  +----- Yaukthwa       
     +--+                     
        |  +-- Lens_contradens **
        +--+                  
           +-- Lens_eximius **   

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

iTtTbiriSoTroTroSiD
22027811--
  +-- Lens_contradens **
--+                  
  +-- Lens_comptus **   

Dai et al., 2024 [Publ 4389], fig. 3

iTtTbiriSoTroTroSiD
22024511--
  +----- Physunio       
--+                     
  |  +-- Lens_comptus **   
  +--+                  
     +-- Lens_contradens **

Liu et al., 2024 [Publ 4380], fig. 5

iTtTbiriSoTroTroSiD
22023553--
        +-- Physunio       
  +-----+                  
  |     +-- Trapezoideus   
--+                        
  |  +----- Yaukthwa       
  +--+                     
     |  +-- Lens_contradens **
     +--+                  
        +-- Lens_eximius **   

Dai et al., 2023 [Publ 4285], fig. 3-ML

iTtTbiriSoTroTroSiD
22024311--
  +----- Physunio       
--+                     
  |  +-- Lens_contradens **
  +--+                  
     +-- Lens_comptus **   

Wu et al., 2024 [Publ 4350], fig. 5c

iTtTbiriSoTroTroSiD
22023553--
  +----- Physunio           
  |                         
  +----- Yaukthwa_paiensis  
  |                         
  +----- Yaukthwa_inlenensis
--+                         
  +----- Trapezoideus       
  |                         
  +----- Yaukthwa_elongatula
  |                         
  |  +-- Lens_contradens **    
  +--+                      
     +-- Lens_eximius **       

Pfeiffer et al., 2018 [Publ 3827], fig. 1

iTtTbiriSoTroTroSiD
2202522--
  +----- Physunio       
  |                     
--+----- Trapezoideus   
  |                     
  |  +-- Lens_comptus **   
  +--+                  
     +-- Lens_contradens **

Dai et al., 2025 [Publ 4441], fig. 1

iTtTbiriSoTroTroSiD
22024611--
  +----- Physunio       
--+                     
  |  +-- Lens_comptus **   
  +--+                  
     +-- Lens_contradens **

Wu et al., 2024 [Publ 4350], fig. 6-7

iTtTbiriSoTroTroSiD
22023553--
        +-- Physunio       
  +-----+                  
  |     +-- Trapezoideus   
--+                        
  |  +----- Yaukthwa       
  +--+                     
     |  +-- Lens_contradens **
     +--+                  
        +-- Lens_eximius **   

Wu et al., 2024 [Publ 4350], fig. 5a

iTtTbiriSoTroTroSiD
22023553--
        +-- Physunio       
  +-----+                  
  |     +-- Trapezoideus   
--+                        
  |  +----- Yaukthwa       
  +--+                     
     |  +-- Lens_contradens **
     +--+                  
        +-- Lens_eximius **   

Wu et al., 2024 [Publ 4388], fig. 5

iTtTbiriSoTroTroSiD
22023653--
        +-- Physunio       
  +-----+                  
  |     +-- Trapezoideus   
--+                        
  |  +----- Yaukthwa       
  +--+                     
     |  +-- Lens_contradens **
     +--+                  
        +-- Lens_eximius **   

Wu et al., 2024 [Publ 4350], fig. 5b

iTtTbiriSoTroTroSiD
22023553--
  +-------------- Yaukthwa_elongatula
  |                                  
--+  +----------- Trapezoideus       
  |  |                               
  +--+        +-- Physunio           
     |  +-----+                      
     |  |     +-- Yaukthwa_inlenensis
     +--+                            
        |  +----- Yaukthwa_paiensis  
        +--+                         
           |  +-- Lens_contradens **    
           +--+                      
              +-- Lens_eximius **       

Dai et al., 2023 [Publ 4285], fig. 3-BI

iTtTbiriSoTroTroSiD
22024311--
  +----- Physunio       
--+                     
  |  +-- Lens_contradens **
  +--+                  
     +-- Lens_comptus **   

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

iTtTbiriSoTroTroSiD
11019032--
  +----- Physunio    
--+                  
  |  +-- Lens_eximius **
  +--+               
     +-- Yaukthwa    

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

iTtTbiriSoTroTroSiD
11012211--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

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

iTtTbiriSoTroTroSiD
11018711--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

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

iTtTbiriSoTroTroSiD
11015644--
        +-- Trapezoideus   
  +-----+                  
  |     +-- Physunio       
--+                        
  |  +----- Yaukthwa       
  +--+                     
     |  +-- Lens_contradens **
     +--+                  
        +-- Solenaia       

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

iTtTbiriSoTroTroSiD
11014500--
-- Lens_contradens **

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

iTtTbiriSoTroTroSiD
11019032--
  +----- Physunio    
--+                  
  |  +-- Lens_eximius **
  +--+               
     +-- Yaukthwa    

Konopleva et al., 2017 [Publ 3807], fig. 6

iTtTbiriSoTroTroSiD
11013021--
  +-- Lens_micropterus **
--+                   
  +-- Physunio        

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

iTtTbiriSoTroTroSiD
11014600--
-- Lens_contradens **

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

iTtTbiriSoTroTroSiD
11015644--
        +-- Trapezoideus   
  +-----+                  
  |     +-- Physunio       
--+                        
  |  +----- Yaukthwa       
  +--+                     
     |  +-- Lens_contradens **
     +--+                  
        +-- Solenaia       

Jeratthitikul et al., 2024 [Publ 4387], fig. 1

iTtTbiriSoTroTroSiD
11015722--
  +----- Physunio       
--+                     
  |  +-- Lens_contradens **
  +--+                  
     +-- Trapezoideus   

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

iTtTbiriSoTroTroSiD
11014711--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

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

iTtTbiriSoTroTroSiD
11013500--
-- Lens_contradens **

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

iTtTbiriSoTroTroSiD
11015311--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

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

iTtTbiriSoTroTroSiD
11019411--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

Jeratthitikul et al., 2021 [Publ 4219], fig. 2

iTtTbiriSoTroTroSiD
11013522--
  +----- Lens_contradens **
--+                     
  |  +-- Trapezoideus   
  +--+                  
     +-- Physunio       

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

iTtTbiriSoTroTroSiD
11014500--
-- Lens_contradens **

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

iTtTbiriSoTroTroSiD
11014600--
-- Lens_contradens **

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

iTtTbiriSoTroTroSiD
11019011--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

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

iTtTbiriSoTroTroSiD
11015211--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

Jeratthitikul et al., 2021 [Publ 4219], fig. 1

iTtTbiriSoTroTroSiD
11013522--
  +----- Lens_contradens **
--+                     
  |  +-- Trapezoideus   
  +--+                  
     +-- Physunio       

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

iTtTbiriSoTroTroSiD
11013500--
-- Lens_contradens **

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

iTtTbiriSoTroTroSiD
11019011--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

Matos et al., 2024 [Publ 4384], fig. 3

iTtTbiriSoTroTroSiD
11011911--
  +-- Physunio       
--+                  
  +-- Lens_contradens **

Zhang et al., 2024 [Publ 4397], fig. 5

iTtTbiriSoTroTroSiD
11012711--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

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

iTtTbiriSoTroTroSiD
11013921--
  +-- Lens_micropterus **
--+                   
  +-- Physunio        

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

iTtTbiriSoTroTroSiD
110112022--
-- Lens_contradens **

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

iTtTbiriSoTroTroSiD
11019411--
  +-- Lens_contradens **
--+                  
  +-- Physunio       

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

iTtTbiriSoTroTroSiD
11017522--
  +----- Lens_contradens **
--+                     
  |  +-- Trapezoideus   
  +--+                  
     +-- Physunio       

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

iTtTbiriSoTroTroSiD
11017522--
  +----- Lens_contradens **
--+                     
  |  +-- Trapezoideus   
  +--+                  
     +-- Physunio       

Smith, 2021 [Publ 4331], fig. 1

iTtTbiriSoTroTroSiD
11011900--
-- Lens_contradens **

Graf, 2002 [Publ 2651], fig. 2

iTtTbiriSoTroTroSiD
11011700--
-- Lens_contradens **

Matos et al., 2024 [Publ 4383], fig. 3

iTtTbiriSoTroTroSiD
11011911--
  +-- Physunio       
--+                  
  +-- Lens_contradens **
 
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