Bivalvia | Unionida | Unionoidea | Margaritiferidae

subfamily GIBBOSULINAE Bogan, Bolotov, Froufe & Lopes-Lima in Lopes-Lima et al., 2018: cladomics

1 genus

Cladomics pages are experimental — even more so than the rest of this site!

The purpose of the cladomics pages for each genus and family-group level taxon is to provide statistics and references indicating the extent to which published phylogenetic work supports the classification used here.

The Cladomics Summary provides an overview of the number of studies that have included terminal species representing this taxon as well as an indication of how frequently it has been recovered as monophyletic.

Following the summary, the available published clades representing the taxon are listed.

For more information, visit the Cladomics FARQ (frequently asked rhetorical questions) page. Abbreviations for the various statistics are explained on the FARQ!

Cladomics Summary

number of species = 5 (5 total)
number of subtaxa (rank: genus) = 1
analyzed in 81 cladograms in 46 publications
maximum number of terminals in an analysis = 4 (mean = 1.35, median = 1)
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) = 21
maximum number of included terminals = 4, (mean = 2.33 median = 2)
maximum number of subtaxa (genus) = 1 (mean = 1, median = 1)
proportion of cladograms that support basic monophyly = 0.81

Pairwise Cladogram Clustering

The graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.

PCoA Ordination of Gibbosulinae cladogram distances
PCoA Ordination

NJ Tree of Gibbosulinae cladogram distances
NJ Tree

Cladograms

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

** Top Monophyly Cladogram **

iTtTbiriSoTroTroSiD
4 4 2 1 22 11 1 0 
  +----------- Pseudunio_marocanus      
  |                                     
  |        +-- Pseudunio_auricularius__1
--+        |                            
  |  +-----+-- Pseudunio_auricularius__2
  |  |     |                            
  +--+     +-- Pseudunio_homsensis      
     |                                  
     |  +----- Gibbosula **                
     +--+                               
        |  +-- Cumberlandia             
        +--+                            
           +-- Margaritifera

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

** Top Sister-Group Cladogram **

iTtTbiriSoTroTroSiD
3 3 1 1 13 13 1 3 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
3 3 1 1 138 11 1 3 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
3 3 1 1 628 11 1 1 
  +-------- Pseudunio    
--+                      
  |  +----- Gibbosula **    
  +--+                   
     |  +-- Cumberlandia 
     +--+                
        +-- Margaritifera

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

iTtTbiriSoTroTroSiD
3 3 1 1 25 11 1 3 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
3 3 1 1 25 11 1 3 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 94 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 66 10 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Pseudunio    
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Margaritifera

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

iTtTbiriSoTroTroSiD
2 2 0 1 72 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 32 10 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Margaritifera
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Pseudunio

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

iTtTbiriSoTroTroSiD
2 13 0 1 198 11 1 13 
  +----- Gibbosula_rochechouartii **
--+                              
  |  +-- Gibbosula_laosensis **     
  +--+                           
     +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 12 0 1 32 10 1 12 
  +----- Gibbosula_rochechouartii **
--+                              
  |  +-- Gibbosula_laosensis **     
  +--+                           
     +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 66 10 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Margaritifera
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Pseudunio

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

iTtTbiriSoTroTroSiD
2 13 0 1 196 11 1 13 
  +----- Gibbosula_rochechouartii **
--+                              
  |  +-- Gibbosula_laosensis **     
  +--+                           
     +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 94 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 13 0 1 198 11 1 13 
  +----- Gibbosula_rochechouartii **
--+                              
  |  +-- Gibbosula_laosensis **     
  +--+                           
     +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 72 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 27 10 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 8 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
2 2 0 1 66 10 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Pseudunio    
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Margaritifera

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

iTtTbiriSoTroTroSiD
2 2 0 1 67 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 1 1 -- 
  +-- Margaritiferinae
--+                   
  +-- Gibbosula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 75 9 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Margaritifera
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Pseudunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 48 3 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 46 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Margaritiferinae
--+                   
  +-- Gibbosula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 1 1 -- 
  +-- Margaritiferinae
--+                   
  +-- Gibbosula **

Bolotov et al., 2016 [Publ 3732], fig. 2

iTtTbiriSoTroTroSiD
1 1 0 1 7 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 55 0 0 -- 
-- Gibbosula_laosensis **

Vikhrev et al., 2017 [Publ 3810], fig. 4

iTtTbiriSoTroTroSiD
1 1 0 1 9 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Margaritiferinae
--+                   
  +-- Gibbosula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 45 3 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 94 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 137 9 1 -- 
  +----- Cumberlandia 
  |                   
--+  +-- Gibbosula **    
  |  |                
  +--+-- Pseudunio    
     |                
     +-- Margaritifera

Bespalaya et al., 2018 [Publ 3811], fig. 3

iTtTbiriSoTroTroSiD
1 1 0 1 9 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 73 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Bogan et al., 2023 [Publ 4295], fig. 5

iTtTbiriSoTroTroSiD
1 1 0 1 5 0 0 -- 
-- Gibbosula_crassa **

Dai et al., 2026 [Publ 4757], fig. 3

iTtTbiriSoTroTroSiD
1 1 0 1 33 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Bogan et al., 2023 [Publ 4295], fig. 4

iTtTbiriSoTroTroSiD
1 1 0 1 6 0 0 -- 
-- Gibbosula_crassa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 55 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 9 1 -- 
  +-------- Pseudunio    
--+                      
  |  +----- Margaritifera
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Cumberlandia

Chen et al., 2025 [Publ 4663], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 47 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 53 2 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 6 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Bolotov et al., 2022 [Publ 4250], fig. 8

iTtTbiriSoTroTroSiD
1 1 0 1 48 1 1 -- 
-- Gibbosula_laosensis **

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 90 6 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 12 10 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 21 1 1 -- 
  +-- Margaritiferinae
--+                   
  +-- Gibbosula **

Huang et al., 2019 [Publ 3843], fig. S1

iTtTbiriSoTroTroSiD
1 1 0 1 46 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Dai et al., 2024 [Publ 4385], fig. 3-BI

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 41 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 75 9 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Gibbosula **    
  +--+                   
     |  +-- Pseudunio    
     +--+                
        +-- Margaritifera

Chen et al., 2020 [Publ 4170], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 33 0 0 -- 
-- Gibbosula_crassa **

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

iTtTbiriSoTroTroSiD
1 1 0 1 16 1 1 -- 
  +-- Margaritiferinae
--+                   
  +-- Gibbosula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Dai et al., 2024 [Publ 4385], fig. 3-ML

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Sano et al., 2020 [Publ 4138], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 33 2 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 53 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Bogan et al., 2023 [Publ 4295], fig. 3

iTtTbiriSoTroTroSiD
1 1 0 1 30 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 29 9 1 -- 
  +-------- Pseudunio    
--+                      
  |  +----- Margaritifera
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Cumberlandia

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

iTtTbiriSoTroTroSiD
1 1 0 1 50 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 87 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 116 4 1 -- 
  +-- Gibbosula **                  
  |                              
  +-- Margaritifera_dahurica     
  |                              
--+-- Margaritifera_margaritifera
  |                              
  +-- Margaritifera_laevis       
  |                              
  +-- Margaritifera_middendorffi

Zhou et al., 2007 [Publ 2658], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 20 2 1 -- 
-- Gibbosula_rochechouartii **

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

iTtTbiriSoTroTroSiD
1 1 0 1 40 6 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 12 10 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 137 9 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Margaritifera
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Pseudunio

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

iTtTbiriSoTroTroSiD
1 1 0 1 10 10 1 -- 
  +-------- Cumberlandia 
--+                      
  |  +----- Pseudunio    
  +--+                   
     |  +-- Gibbosula **    
     +--+                
        +-- Margaritifera

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

iTtTbiriSoTroTroSiD
1 1 0 1 13 2 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 85 5 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 52 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

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

iTtTbiriSoTroTroSiD
1 1 0 1 64 1 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Huang et al., 2019 [Publ 3843], fig. 1

iTtTbiriSoTroTroSiD
1 1 0 1 44 1 1 -- 
  +-- Margaritiferinae
--+                   
  +-- Gibbosula **

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

iTtTbiriSoTroTroSiD
1 1 0 1 90 6 1 -- 
  +-- Gibbosula **       
--+                   
  +-- Margaritiferinae

Bogan et al., 2023 [Publ 4295], fig. 6

iTtTbiriSoTroTroSiD
1 1 0 1 12 0 0 -- 
-- Gibbosula_crassa **