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Page last updated
Thu 01 May 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!

This page documents data for Pfeiffer & Graf (2015), such as genera and species attributed to that publication (including known misspellings, misidentifications, and other unavailable names) as well as taxonomic opinions, if available. Taxonomic opinions refer to uses of genera and species as valid and invalid in that work.

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Publication information

Pfeiffer, J.M. & D.L. Graf. 2015. Evolution of bilaterally asymmetrical larvae in freshwater mussels (Bivalvia: Unionoida: Unionidae). Zoological Journal of the Linnean Society 175(2): 307-318.

Click here for online access.

Cladogram Figures

(click the figure number to see the cladogram)

fig. 2

Attributed Species Names

Pseudodon vondembuschianus Graf, 2002
‘Lea’ of Authors.
Graf, 2002, J. Moll. Studies 68: 66.
Graf & Cummings, 2006, Zool. J. Linn. Soc. 148: 348.
Hoeh et al., 2009, Malacologia 51(2): 310.
Whelan et al., 2011, MPE 61: 506. [in part]
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310. [in part]
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Pseudodon aff. vondembuschianus sp. 2’]. [in part]
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207. [in part]
Bolotov et al., 2018, Sci. Repts. 8: Table S1 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
Huang et al., 2019, MPE 130: Table S4 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
(reference to Margaritana vondembuschiana Lea, 1840)

Velesunio ambiguous Graf & Cummings, 2006
‘Philippi’ Graf & Cummings, 2006, Zool. J. Linn. Soc. 148: 348, table 2.
‘Philippi’ Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): [4].
(error for Unio ambiguus Philippi, 1847)

Radiatula bonneaudii Whelan et al., 2011
‘Eydoux’ of Authors.
Whelan et al., 2011, MPE 61: 506.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): [4].
Bolotov et al., 2017, Sci. Repts. 7: 10, Table 3 [as ‘sp. 1’].
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Bolotov et al., 2017, Sci. Repts. 7: [5].
Bolotov et al., 2018, Sci. Repts. 8: Table S1.
Pfeiffer et al., 2018, J. Moll. Studies 84(4): [4]. [in part]
Jeratthitikul et al., 2019, Tropical Nat. Hist. 19(1): 25.
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘Radiatula bonneaudii sp. 1’].
Huang et al., 2019, MPE 130: Table S4.
Ortiz-Sepulveda et al., 2020, MPE 148: Table S1.
(misidentified reference to Unio bonneaudii Eydoux, 1838)

Lamellidens corrianus Whelan et al., 2011
‘Lea’ of Authors.
Whelan et al., 2011, MPE 61: 506.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Lamellidens corrianus sp. 1].
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Pfeiffer et al., 2018, J. Moll. Studies 84(4): [3].
Pfeiffer et al., 2019, MPE 137: 117.
Lopes-Lima et al., 2020, MPE 146: Table 1.
(misidentified reference to Unio corrianus Lea, 1834)
Type(s): SEQUENCED SPECIMEN UMMZ_304642, Ayeryarwaddy River, mud bank on E side of river, Mandalay Division, Burma. 21.48187, 95.30501.

Lamellidens generosus Whelan et al., 2011
‘Gould’ of Authors.
Whelan et al., 2011, MPE 61: 506.
Pfeiffer & Graf, 2013, J. Moll. Studies 79: 251.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Araujo et al., 2017, Zool. J. Linn. Soc. 179(2): 280.
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Pfeiffer et al., 2018, J. Moll. Studies 84(4): [3].
Pfeiffer et al., 2019, MPE 137: 117.
Wu et al., 2021, J. Lake Sci. 33(6): attached Table 1.
(misidentified reference to Unio generosus Gould, 1847)

Pseudodon inoscularis Pfeiffer & Graf, 2013
‘Gould’ of Authors.
Pfeiffer & Graf, 2013, J. Moll. Studies 79: 251.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Pseudodon cf. inoscularis’].
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘ “Monodontina” cf. inoscularis sp. 2’].
Huang et al., 2019, MPE 130: Table S4 [as ‘ “Monodontina” cf. inoscularis sp. 2’].
Lopes-Lima et al., 2020, MPE 146: Table 1 [as ‘ “Monodontina” cf. inoscularis sp. 2’].
(misidentified reference to Anodon inoscularis Gould, 1844)

Trapezoideus exolescens Pfeiffer & Graf, 2013
‘Gould’ of Authors.
Brandt, 1974, Arch. Moll. 105: 299.
Pfeiffer & Graf, 2013, J. Moll. Studies 79: 251.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): [4].
Lopes-Lima et al., 2017, MPE 106: 177.
(misidentified reference to Unio exolescens Gould, 1843)

Pseudodon cambodjensis Pfeiffer & Graf, 2013
‘Petit’ of Authors.
Pfeiffer & Graf, 2013, J. Moll. Studies 79: 251.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Zieritz et al., 2016, Sci. Total Environ. 571: 1071.
Bolotov et al., 2017, Sci. Repts. 7: Table S1.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Monodontina cambodjensis’].
Lopes-Lima et al., 2017, MPE 106: 177.
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Bolotov et al., 2018, Sci. Repts. 8: Table S1 [as ‘Monodontina cambodjensis’].
Razak et al., 2018, Conserv. Genet. Resour. 11: 459.
Huang et al., 2019, MPE 130: Table S4 [as ‘Monodontina cambodjensis’].
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘Monodontina cambodjensis’].
Bolotov et al., 2020, Sci. Repts. 10: Table S1 [as ‘Monodontina cambodiensis’ (sic)].
Konopleva et al., 2021, Sci. Repts. 11: Table S1 [as ‘Monodontina cambodiensis’ (sic)].
Jeratthitikul et al., 2021, Eur. J. Taxonomy 775: 123 [as ‘Monodontina cambodiensis’ (sic)].
Pfeiffer et al., 2021, Invert. Syst. 35: 453 [as ‘Monodontina cambodjensis’].
Wu et al., 2021, ZooKeys 1054: 87 [as ‘Monodontina cambodjensis’].
Wu et al., 2021, J. Lake Sci. 33(6): attached Table 1.
Konopleva et al., 2023, Diversity 15(1), 10: Table S2 [as ‘Monodontina cambodjensis’].
Dai et al., 2023, Invert. Syst. 37(2): Table 1 [as ‘Monodontina cambodjensis’].
Dai et al., 2024, Zoosyst. Evol. 100(4): Table 2.
(misidentified reference to Monocondylea cambodjensis Petit de la Saussaye, 1865)
Type(s): SEQUENCED SPECIMEN UMMZ_304350, Pursat River, just downstream (north) of Highway 5 bridge, Pursat City, Cambodia.

Pseudodon vondembuschianus Pfeiffer & Graf, 2015
‘Lea’ of Authors.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310. [in part]
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Pseudodon aff. vondembuschianus sp. 2’]. [in part]
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207. [in part]
Bolotov et al., 2018, Sci. Repts. 8: Table S1 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
Huang et al., 2019, MPE 130: Table S4 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘Monodontina aff. vondembuschiana sp. 2’]. [in part]
(reference to Margaritana vondembuschiana Lea, 1840)

Scabies crispata Pfeiffer & Graf, 2015
‘Gould’ of Authors.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): [4].
Ng et al., 2016, PLoS One 11(8), e0161130: [5].
Bolotov et al., 2017, Sci. Repts. 7: [6], Table S1 [as ‘Scabies aff. crispata’].
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
(misidentified reference to Unio crispata Gould, 1843)
Type(s): SEQUENCED SPECIMEN UMMZ_304646, Beung Dang Teuk Leach Community Fishery Pursat Province, Cambodia.

Pilsbryoconcha exilis Pfeiffer & Graf, 2015
‘Lea’ of Authors.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Ng et al., 2016, PLoS One 11(8), e0161130: [5]. [in part]
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘P. aff. exilis sp. 1’].
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘P. aff. exilis sp. 1’].
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Bolotov et al., 2018, Sci. Repts. 8: Table S1 [as ‘P. aff. exilis sp. 1’].
Huang et al., 2019, MPE 130: Table S4 [as ‘P. aff. exilis sp. 1’].
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘P. aff. exilis sp. 1’].
Lopes-Lima et al., 2020, MPE 146: Table 1 [as ‘P. aff. exilis sp. 1’].
Zieritz et al., 2020, Biodivers. Conserv. 29: 2244.
Wu et al., 2021, ZooKeys 1054: 87.
Wu et al., 2021, J. Lake Sci. 33(6): attached Table 1.
Wu et al., 2021, J. Lake Sci. 33(6): attached Table 1.
Konopleva et al., 2023, Diversity 15(1), 10: Table S2 [as ‘P. aff. exilis’].
Dai et al., 2023, Invert. Syst. 37(2): Tables 1-2. [in part]
(misidentified reference to Anodonta exilis Lea, 1838)
Type(s): SEQUENCED SPECIMEN UMMZ_304647, Beung Dang Teuk Leach Community Fishery Pursat Province, Cambodia.

Ensidens sagittarius Pfeiffer & Graf, 2015
‘Lea’ of Authors.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Lopes-Lima et al., 2017, MPE 106: 177.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Ensidens aff. sagittarius sp. 1].
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Ensidens aff. sagittarius sp. 1].
Bolotov et al., 2018, Sci. Repts. 8: Table S1 [as ‘Ensidens aff. sagittarius sp. 1].
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘E. aff. sagittarius sp. 1].
Huang et al., 2019, MPE 130: Table S4 [as ‘Ensidens aff. sagittarius sp. 1].
Zieritz et al., 2020, Biodivers. Conserv. 29: 2242.
Lopes-Lima et al., 2020, MPE 146: Table 1.
Wu et al., 2021, J. Lake Sci. 33(6): attached Table 1.
(misidentified reference to Unio sagittarius Lea, 1856)

Contradens contradens Pfeiffer & Graf, 2015
‘Lea’ of Authors.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Contradens aff. contradens sp. 1’].
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Contradens aff. contradens sp. 1’].
Bolotov et al., 2018, Sci. Repts. 8: Table S1 [as ‘Contradens aff. contradens sp. 1’].
Konopleva et al., 2019, Ecologica Montenegrina 24: Appendix 1 [as ‘Contradens aff. contradens sp. 1’].
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘Contradens aff. contradens sp. 1’].
Huang et al., 2019, MPE 130: Table S4 [as ‘Contradens aff. contradens sp. 1’].
Bolotov et al., 2020, Sci. Repts. 10: Table S1 [as ‘Contradens aff. contradens sp. 1’].
Konopleva et al., 2023, Diversity 15(1), 10: Table S2 [as ‘Lens aff. contradens’].
(misidentified reference to Unio contradens Lea, 1838)

Pseudodon cumingii Pfeiffer & Graf, 2015
‘Lea’ of Authors.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Lopes-Lima et al., 2017, MPE 106: 177.
Bolotov et al., 2017, Sci. Repts. 7: Table S1.
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘”Bineurus” cumingii’].
Huang et al., 2019, MPE 130: Table S4 [as ‘”Bineurus” cumingii’].
Lopes-Lima et al., 2020, MPE 146: Table 1.
(misidentified reference to Anodonta cumingii Lea, 1851)
Type(s): SPECIMEN UMMZ_304648, Nam Phiat, Phonethong, ca. 2 km from confluence, Bolikhamxai Province, Laos. 18.08389, 104.97806.

Pseudodon mouhotii Pfeiffer & Graf, 2015
‘Lea’ of Authors.
Pfeiffer & Graf, 2015, Zool. J. Linn. Soc. 175(2): 310.
Lopes-Lima et al., 2017, MPE 106: 177.
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Pseudodon aff. mouhotii sp. 3’].
Bolotov et al., 2017, Sci. Repts. 7: Table S1 [as ‘Bineurus aff. mouhotii sp. 3’].
Konopleva et al., 2017, Systemat. Biodiv. 15(3): 207.
Bolotov et al., 2018, Sci. Repts. 8: Table S1 [as ‘Bineurus aff. mouhotii sp. 3’].
Konopleva et al., 2019, Sci. Repts. 9: Table S1 [as ‘Bineurus aff. mouhotii sp. 3’].
Huang et al., 2019, MPE 130: Table S4 [as ‘Bineurus aff. mouhotii sp. 3’].
(misidentified reference to Monocondyloea mouhotii Lea, 1863)

Taxonomic Opinions

Amblema plicata (Say, 1817)

Castalia ambigua Lamarck, 1819

Chambardia wahlbergi (Krauss, 1848)

Coelatura aegyptiaca (Cailliaud, 1827)

Contradens sp.
+ Physunio modelli Brandt, 1974

Ensidens sagittarius (Lea, 1856)

Etheria elliptica Lamarck, 1807

Gonidea angulata (Lea, 1838)

Hyridella australis (Lamarck, 1819)

Lamellidens corrianus (Lea, 1834)

Lamellidens generosus (Gould, 1847)

Lamproscapha ensiformis (Spix & Wagner, 1827)

Lamprotula caveata (Heude, 1877)

Lampsilis cardium Rafinesque, 1820

Lasmigona costata
+ Lasmigona compressa Graf & Ó Foighil, 2000

Margaritifera margaritifera (Linnaeus, 1758)

Margaritifera monodonta (Say, 1829)

Neotrigonia margaritacea (Lamarck, 1804)

Oxynaia pugio (Benson, 1862)

Parreysia olivacea Prashad, 1930

Parreysia tavoyensis (Gould, 1843)

Pilsbryoconcha exilis (Lea, 1838)

Potamilus alatus (Say, 1817)

Potomida littoralis (Cuvier, 1798)

Pronodularia japanensis (Lea, 1859)

Pseudodon cambodjensis (Petit, 1865)

Pseudodon cumingii (Lea, 1850)

Pseudodon inoscularis (Gould, 1844)

Pseudodon mouhotii (Lea, 1863)

Pseudodon vondembuschianus (Lea, 1840)

Pyganodon grandis (Say, 1829)

Quadrula quadrula (Rafinesque, 1820)

Radiatula bonneaudii (Eydoux, 1838)

Scabies crispata (Gould, 1843)

Trapezoideus exolescens (Gould, 1843)

Triplodon corrugatus (Lamarck, 1819)

Truncilla truncata Rafinesque, 1820

Unio pictorum (Linnaeus, 1758)

Velesunio ambiguous [sic] (Philippi, 1847)

Cladogram Figure 2

  +-------------------------------------- Neotrigonia_margaritacea   
  |                                                                  
  |                             +-------- Velesunio_ambiguus         
  |  +--------------------------+                                    
  |  |                          |  +----- Hyridella_australis        
--+  |                          +--+                                 
  |  |                             |  +-- Triplodon_corrugatus       
  |  |                             +--+                              
  |  |                                +-- Castalia_ambigua           
  |  |                                                               
  +--+                             +----- Etheria_elliptica          
     |  +--------------------------+                                 
     |  |                          |  +-- Lamproscapha_ensiformis    
     |  |                          +--+                              
     |  |                             +-- Chambardia_wahlbergi       
     |  |                                                            
     |  |                             +-- Margaritifera_margaritifera
     |  |  +--------------------------+                              
     +--+  |                          +-- Cumberlandia_monodonta     
        |  |                                                         
        |  |                          +-- Trapezidens_dolichorhynchus
        |  |              +-----------+                              
        |  |              |           +-- Lamellidens_savadiensis    
        |  |              |                                          
        |  |  +-----------+           +-- Leoparreysia_olivacea      
        +--+  |           |  +--------+                              
           |  |           |  |        +-- Leoparreysia_tavoyensis    
           |  |           +--+                                       
           |  |              |  +-------- Coelatura_aegyptiaca       
           |  |              +--+                                    
           |  |                 |  +----- Indochinella_pugio         
           |  |                 +--+                                 
           |  |                    |  +-- Radiatula_chaudhurii       
           |  |                    +--+                              
           +--+                       +-- Scabies_mandarinus         
              |                                                      
              |                    +----- Unio_pictorum              
              |  +-----------------+                                 
              |  |                 |  +-- Pyganodon_grandis          
              |  |                 +--+                              
              |  |                    +-- Lasmigona_costata          
              |  |                                                   
              |  |           +----------- Quadrula_quadrula          
              |  |  +--------+                                       
              +--+  |        |  +-------- Amblema_plicata            
                 |  |        +--+                                    
                 |  |           |  +----- Potamilus_alatus           
                 |  |           +--+                                 
                 |  |              |  +-- Lampsilis_cardium          
                 |  |              +--+                              
                 |  |                 +-- Truncilla_truncata         
                 +--+                                                
                    |           +-------- Ensidens_telus             
                    |  +--------+                                    
                    |  |        |  +----- Lens_micropterus           
                    |  |        +--+                                 
                    |  |           |  +-- Physunio                   
                    |  |           +--+                              
                    |  |              +-- Physunio_modelli           
                    +--+                                             
                       |        +-------- Gonidea_angulata           
                       |  +-----+                                    
                       |  |     |  +----- Potomida_littoralis        
                       |  |     +--+                                 
                       |  |        |  +-- Pronodularia_japanensis    
                       +--+        +--+                              
                          |           +-- Lamprotula_caveata         
                          |                                          
                          |  +----------- Pilsbryoconcha_carinifera  
                          |  |                                       
                          +--+        +-- Pseudodon_vondembuschianus 
                             |  +-----+                              
                             |  |     +-- Pseudodon_vagulus          
                             +--+                                    
                                |  +----- Sundadontina_tumida        
                                +--+                                 
                                   |  +-- Nyeinchanconcha_nyeinchani 
                                   +--+                              
                                      +-- Bineurus_exilis

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