MUSSELpdb
Taxonomy
Species & Genus Richness of
freshwater mussels
by Taxonomy
Species & Genus Richness of
freshwater bivalves
by Family
Geography
Species Richness & Endemism of
freshwater mussels
by Geography
Data last updated
Thu 19 March 2026
Publication Sano et al. (2020): cladograms
Complete Citation
Sano, I., T. Saito, J.-I. Miyazaki, A. Shirai, T. Uechi et al. 2020. Evolutionary history and diversity of unionoid mussels (Mollusca: Bivalvia) in the JApanese Archipelago. Plankton Benthos Res 15(2): 97-111. Available electronically.
Cladogram Figures
(click the figure number to see the cladogram)
Cladogram Figure 1
+-------------------- Neotrigonia_lamarckii
|
| +----- Gibbosula_rochechouartii
| +-----------+
| | | +-- Margaritifera_middendorffi
| | +--+
| | +-- Margaritifera_laevis
| |
| | +-------- Sinosolenaia_iridinea
--+ | |
| | +--+ +----- Inversidens_brandtii
| | | | |
| | | +--+ +-- Obovalis_omiensis__1
| | | | |
| | | +--+-- Obovalis_omiensis__2
| | +--+ |
| | | | +-- Ptychorhynchus_pfisteri
| | | |
| | | | +-- Lamprotula_caveata
+--+ | | +--+
| | | | +-- Pronodularia_japanensis
| | +-----+
| | | +-- Sinohyriopsis_schlegelii__1
| | | |
| | +--+-- Sinohyriopsis_schlegelii__2
| | |
| | +-- Sinohyriopsis_cumingii
| |
| | +-------- Lanceolaria_grayii__1
| | |
| | +-------- Lanceolaria_grayii__2
| | |
| | +-------- Cristaria_plicata__1
| | |
| | +-------- Cristaria_plicata__2
+--+ |
| +-------- Sinanodonta_lauta
| |
| +-------- Beringiana_japonica
| |
| +-------- Beringiana_fukuharai__1
| |
| +-------- Beringiana_beringiana
| |
| +-------- Beringiana_fukuharai__2
| |
| +-------- Beringiana_fukuharai__3
| |
| +-------- Buldowskia_iwakawai__1
| |
| +-------- Buldowskia_iwakawai__2
| |
| +-------- Sinanodonta_tumens__1
| |
| +-------- Sinanodonta_tumens__2
| |
+-----+-------- Pletholophus_reinianus
|
+-------- Anemina_harlandi
|
+-------- Lanceolaria_lanceolata
|
+-------- Cristaria_plicata__3
|
+-------- Lanceolaria_gladiola
|
+-------- Sinanodonta_angula
|
+-------- Lepidodesma_languilati
|
| +-- Scriptolamprotula_tortuosa
+-----+
| +-- Acuticosta_chinensis
|
| +----- Cuneopsis_celtiformis
| |
+--+ +-- Nodularia_nipponensis__1
| | |
| +--+-- Nodularia_douglasiae
| |
| +-- Nodularia_nipponensis__2
|
| +----- Schistodesmus_xinyuensis
| |
| | +-- Inversiunio_reinianus__1
+--+ |
| +-- Inversiunio_reinianus__2
| |
| +-- Inversiunio_jokohamensis__1
+--+
+-- Inversiunio_jokohamensis__2
|
+-- Inversiunio_jokohamensis__3
|
+-- Inversiunio_jokohamensis__4
Cladogram Figure 2
+-- Neotrigonia_margaritacea
+--------------------------------------------------+
| +-- Neotrigonia_lamarckii
|
| +----- Triplodon_corrugatus
| +--------------------------------------------+
| | | +-- Hyridella_australis
| | +--+
--+ | +-- Velesunio_ambiguus
| |
| | +----------- Mutela_hargeri
| | |
| | +-----------------------------------+ +-- Aspatharia_pfeifferiana
| | | | +-----+
| | | | | +-- Chambardia_wahlbergi
+--+ | +--+
| | | +----- Etheria_elliptica
| | +--+
| | | +-- Tamsiella_schroeteriana
| | +--+
| | +-- Lamproscapha_ensiformis
| |
| | +-------------------- Cumberlandia_monodonta
| | |
| | | +-- Gibbosula_rochechouartii
| | | +--+
+--+ +-----------------------+ | +-- Gibbosula_laosensis
| | | +-----------+
| | | | | +-- Pseudunio_auricularius
| | | | +--+
| | | | +-- Pseudunio_marocanus
| | +--+
| | | +-- Margaritifera_dahurica
| | | +-----------+
| | | | +-- Margaritifera_margaritifera
| | +--+
| | | +----------- Margaritifera_falcata
| | +--+
| | | +-------- Margaritifera_middendorffi
| | +--+
+--+ | +----- Margaritifera_laevis
| +--+
| | +-- Margaritifera_hembeli
| +--+
| +-- Margaritifera_marrianae
|
| +----------------------------------------- Scabies_crispata
| |
| | +----------------- Lampsilis_cardium
| | |
| | | +----- Sinohyriopsis_cumingii__2
| | +--------------------+ +--------+
| | | | | | +-- Sinohyriopsis_schlegelii
| | | | | +--+
| | | +--+ +-- Sinohyriopsis_cumingii__1
+--+ | |
| | | +----------- Inversidens_brandtii
| | | |
| | +--+ +-- Ptychorhynchus_pfisteri
| | | +-----+
| | | | +-- Obovalis_omiensis
| | +--+
| | | +----- Lamprotula_leaii
| | +--+
+--+ | +-- Pronodularia_japanensis
| +--+
| +-- Potomida_littoralis
|
| +-- Anemina_harlandi
| +--------------------------------+
| | +-- Buldowskia_iwakawai
| |
| | +-------------- Pletholophus_reinianus
| | |
| | | +----- Beringiana_fukuharai
| | +-----------------+ +-----+
| | | | | | +-- Beringiana_japonica
+--+ | | | +--+
| | +--+ +-- Beringiana_beringiana
| | |
| | | +-------- Sinanodonta_lucida
| | +--+
| | | +----- Sinanodonta_tumens
| | +--+
| | | +-- Sinanodonta_lauta
+--+ +--+
| +-- Sinanodonta_woodiana
|
| +----------------------------- Cristaria_plicata
| |
| | +-- Anodonta_cygnea
| | +--------+
| | | +-- Anodonta_anatina
| | |
| | +--------------+ +----- Alasmidonta_varicosa
+--+ | | +--+
| | | | | +-- Prolasmidonta_heterodon
| | | | +--+
| | +--+ +-- Platynaias_compressa
| | |
| | | +----- Pyganodon_grandis
| | +--+
+--+ | +-- Utterbackia_imbecillis
| +--+
| +-- Utterbackia_peninsularis
|
| +----- Lanceolaria_grayii__1
| +-----------------+
| | | +-- Lanceolaria_lanceolata
| | +--+
| | +-- Lanceolaria_grayii__2
+--+
| +-------------------- Lepidodesma_languilati
| |
| | +----- Scriptolamprotula_tortuosa
| | +-----------+
+--+ | | +-- Scriptolamprotula_tientsinensis
| | +--+
| | +-- Scriptolamprotula_coreana
| |
+--+ +----- Unio_nanus
| +--------+
| | | +-- Unio_delphinus
| | +--+
| | +-- Unio_pictorum
| |
+--+ +-------- Inversiunio_jokohamensis
| +--+
| | | +----- Inversiunio_yanagawensis__2
| | +--+
| | | +-- Inversiunio_reinianus
| | +--+
+--+ +-- Inversiunio_yanagawensis__1
|
| +-- Tchangsinaia_piscicula
| +-----+
| | +-- Schistodesmus_lampreyanus
+--+
| +----- Nodularia_nipponensis__1
+--+
| +-- Nodularia_nipponensis__2
+--+
+-- Nodularia_douglasiae
Cladogram Figure 3
+-- Neotrigonia_lamarckii
+-----------------------------------------------------+
| +-- Neotrigonia_margaritacea
|
| +----- Triplodon_corrugatus
| +-----------------------------------------------+
| | | +-- Hyridella_australis
| | +--+
--+ | +-- Velesunio_ambiguus
| |
| | +----------- Mutela_hargeri
| | |
| | +--------------------------------------+ +-- Aspatharia_pfeifferiana
| | | | +-----+
| | | | | +-- Chambardia_wahlbergi
+--+ | +--+
| | | +----- Etheria_elliptica
| | +--+
| | | +-- Tamsiella_schroeteriana
| | +--+
| | +-- Lamproscapha_ensiformis
| |
| | +----------------------- Cumberlandia_monodonta
| | |
| | | +-- Pseudunio_auricularius
| | +-----------------------+ +-----------------+
+--+ | | | +-- Pseudunio_marocanus
| | | |
| | +--+ +-- Gibbosula_laosensis
| | | +--------------+
| | | | +-- Gibbosula_rochechouartii
| | | |
| | +--+ +-- Margaritifera_margaritifera
| | | +-----------+
| | | | +-- Margaritifera_dahurica
| | +--+
| | | +----------- Margaritifera_falcata
| | +--+
| | | +-------- Margaritifera_middendorffi
| | +--+
+--+ | +----- Margaritifera_laevis
| +--+
| | +-- Margaritifera_hembeli
| +--+
| +-- Margaritifera_marrianae
|
| +-------------------------------------------- Scabies_crispata
| |
| | +----- Sinohyriopsis_cumingii__2
| | +--------+
| | | | +-- Sinohyriopsis_schlegelii
| | | +--+
| | | +-- Sinohyriopsis_cumingii__1
| | +--------------------------+
| | | | +----------- Lampsilis_cardium
+--+ | | |
| | | | +----- Lamprotula_leaii
| | +--+ +--+
| | | | | +-- Potomida_littoralis
| | | | +--+
| | +--+ +-- Pronodularia_japanensis
| | |
| | | +----- Inversidens_brandtii
| | +--+
| | | +-- Obovalis_omiensis
+--+ +--+
| +-- Ptychorhynchus_pfisteri
|
| +-- Anemina_harlandi
| +-----------------------------------+
| | +-- Buldowskia_iwakawai
| |
| | +-------- Pletholophus_reinianus
| | +--+
| | | | +----- Beringiana_fukuharai
| | | +--+
| | | | +-- Beringiana_japonica
+--+ +-----------------------+ +--+
| | | +-- Beringiana_beringiana
| | |
| | | +-------- Sinanodonta_lucida
| | +--+
| | | +----- Sinanodonta_tumens
| | +--+
| | | +-- Sinanodonta_woodiana
| | +--+
+--+ +-- Sinanodonta_lauta
|
| +-------------------------------- Cristaria_plicata
| |
| | +-- Anodonta_anatina
| | +--------+
| | | +-- Anodonta_cygnea
| | |
| | +-----------------+ +----- Pyganodon_grandis
+--+ | | +--+
| | | | | +-- Utterbackia_imbecillis
| | | | +--+
| | +--+ +-- Utterbackia_peninsularis
| | |
| | | +----- Alasmidonta_varicosa
| | +--+
+--+ | +-- Platynaias_compressa
| +--+
| +-- Prolasmidonta_heterodon
|
| +----- Lanceolaria_grayii__1
| +--------------------+
| | | +-- Lanceolaria_lanceolata
| | +--+
| | +-- Lanceolaria_grayii__2
+--+
| +----------------------- Lepidodesma_languilati
| |
| | +----- Scriptolamprotula_tortuosa
| | +--------------+
+--+ | | +-- Scriptolamprotula_tientsinensis
| | +--+
| | +-- Scriptolamprotula_coreana
| |
+--+ +----- Unio_nanus
| +-----------+
| | | +-- Unio_delphinus
| | +--+
| | +-- Unio_pictorum
| |
+--+ +-------- Inversiunio_jokohamensis
| +-----+
| | | +----- Inversiunio_yanagawensis__2
| | +--+
| | | +-- Inversiunio_reinianus
| | +--+
+--+ +-- Inversiunio_yanagawensis__1
|
| +-- Schistodesmus_lampreyanus
| +--------+
| | +-- Tchangsinaia_piscicula
+--+
| +-------- Nodularia_nipponensis__1
+--+
| +----- Nodularia_nipponensis__2
+--+
| +-- Nodularia_nipponensis__3
+--+
+-- Nodularia_douglasiae
Cladogram Figure 4
+-- Neotrigonia_lamarckii
+-----------------------------------------------------+
| +-- Neotrigonia_margaritacea
|
| +----- Triplodon_corrugatus
| +-----------------------------------------------+
| | | +-- Hyridella_australis
| | +--+
--+ | +-- Velesunio_ambiguus
| |
| | +----------- Mutela_hargeri
| | |
| | +--------------------------------------+ +-- Aspatharia_pfeifferiana
| | | | +-----+
| | | | | +-- Chambardia_wahlbergi
+--+ | +--+
| | | +----- Etheria_elliptica
| | +--+
| | | +-- Tamsiella_schroeteriana
| | +--+
| | +-- Lamproscapha_ensiformis
| |
| | +-------------------- Cumberlandia_monodonta
| | |
| | | +-- Pseudunio_auricularius
| | +--------------------------+ +--------------+
+--+ | | | +-- Pseudunio_marocanus
| | | |
| | +--+ +-- Gibbosula_rochechouartii
| | | +-----------+
| | | | +-- Gibbosula_laosensis
| | | |
| | +--+ +-- Margaritifera_margaritifera
| | | +--------+
| | | | +-- Margaritifera_dahurica
| | +--+
| | | +-------- Margaritifera_falcata
| | | |
| | +--+ +-- Margaritifera_laevis
| | | +--+
+--+ | | +-- Margaritifera_middendorffi
| +--+
| | +-- Margaritifera_hembeli
| +--+
| +-- Margaritifera_marrianae
|
| +-------------------------------------------- Scabies_crispata
| |
| | +----------------- Lampsilis_cardium
| | |
| | | +----- Sinohyriopsis_cumingii__2
| | +-----------------------+ +--------+
| | | | | | +-- Sinohyriopsis_schlegelii
| | | | | +--+
| | | +--+ +-- Sinohyriopsis_cumingii__1
+--+ | |
| | | +----------- Inversidens_brandtii
| | | |
| | +--+ +-- Obovalis_omiensis
| | | +-----+
| | | | +-- Ptychorhynchus_pfisteri
| | +--+
| | | +----- Lamprotula_leaii
| | +--+
+--+ | +-- Pronodularia_japanensis
| +--+
| +-- Potomida_littoralis
|
| +-- Anemina_harlandi
| +-----------------------------------+
| | +-- Buldowskia_iwakawai
| |
| | +-------- Sinanodonta_lucida
| | +--+
| | | | +----- Sinanodonta_tumens
| | | +--+
| | | | +-- Sinanodonta_woodiana
+--+ +-----------------------+ +--+
| | | +-- Sinanodonta_lauta
| | |
| | | +-------- Pletholophus_reinianus
| | +--+
| | | +----- Beringiana_fukuharai
| | +--+
| | | +-- Beringiana_japonica
| | +--+
+--+ +-- Beringiana_beringiana
|
| +-------------------------------- Cristaria_plicata
| |
| | +-- Anodonta_cygnea
| | +--------+
| | | +-- Anodonta_anatina
| | |
| | +-----------------+ +----- Alasmidonta_varicosa
+--+ | | +--+
| | | | | +-- Prolasmidonta_heterodon
| | | | +--+
| | +--+ +-- Platynaias_compressa
| | |
| | | +----- Pyganodon_grandis
| | +--+
+--+ | +-- Utterbackia_imbecillis
| +--+
| +-- Utterbackia_peninsularis
|
| +-------------------------- Lepidodesma_languilati
| |
| | +----- Lanceolaria_grayii__1
| | +-----------------+
+--+ | | +-- Lanceolaria_lanceolata
| | +--+
| | +-- Lanceolaria_grayii__2
| |
+--+ +----- Scriptolamprotula_tortuosa
| +--------------+
| | | +-- Scriptolamprotula_tientsinensis
| | +--+
| | +-- Scriptolamprotula_coreana
| |
+--+ +----- Unio_nanus
| +-----------+
| | | +-- Unio_delphinus
| | +--+
| | +-- Unio_pictorum
| |
+--+ +-------- Nodularia_nipponensis__3
| +-----+
| | | +----- Nodularia_nipponensis__1
| | +--+
| | | +-- Nodularia_douglasiae
| | +--+
+--+ +-- Nodularia_nipponensis__2
|
| +-- Schistodesmus_lampreyanus
| +--------+
| | +-- Tchangsinaia_piscicula
+--+
| +-------- Inversiunio_jokohamensis
+--+
| +----- Inversiunio_yanagawensis__2
+--+
| +-- Inversiunio_reinianus
+--+
+-- Inversiunio_yanagawensis__1
