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
Mon 29 June 2026
Bivalvia | Unionida | Unionoidea | Margaritiferidae | Gibbosulinae
genus Gibbosula Simpson, 1900: cladomics
5 Recent valid species (5 total species)
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)
analyzed in 85 cladograms in 50 publications
maximum number of terminals in an analysis = 4 (mean = 1.38, median = 1)
Basic Monophyly
number of cladograms that test basic monophyly (i.e., more than 1 included species) = 23
maximum number of included terminals = 4, (mean = 2.39 median = 2)
proportion of cladograms that support basic monophyly = 0.83
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.
Cladograms
Chen et al., 2025 [Publ 4455], fig. 1
** Top Monophyly Cladogram ** ** Top Sister-Group Cladogram **
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 4 | 4 | 2 | 4 | 22 | 0 | 0 | ** |
+-- Cumberlandia
+-----+
| +-- Margaritifera
|
--+ +-- Gibbosula_laosensis **
| +--+
| | +-- Gibbosula_sinensis **
+--+
| +-- Gibbosula_crassa **
+--+
+-- Gibbosula_rochechouartii **
Chen et al., 2026 [Publ 4827], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 4 | 4 | 2 | 4 | 8 | 0 | 0 | 2.0 |
+----- Scriptolamprotula
+-----+
| | +-- Huizhou
| +--+
--+ +-- Globunio
|
| +-------- Gibbosula_crassa **
+--+
| +----- Gibbosula_rochechouartii **
+--+
| +-- Gibbosula_laosensis **
+--+
+-- Gibbosula_sinensis **
Lopes-Lima et al., 2018 [Publ 3829], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 3 | 3 | 1 | 3 | 25 | 0 | 0 | 3.0 |
+----- Gibbosula_rochechouartii **
+--+
| | +-- Gibbosula_crassa **
| +--+
--+ +-- Gibbosula_laosensis **
|
| +----- Margaritifera
+--+
| +-- Cumberlandia
+--+
+-- Pseudunio
Lopes-Lima et al., 2018 [Publ 3829], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 3 | 3 | 1 | 3 | 25 | 0 | 0 | 3.0 |
+----- Gibbosula_rochechouartii **
+--+
| | +-- Gibbosula_laosensis **
| +--+
--+ +-- Gibbosula_crassa **
|
| +----- Margaritifera
+--+
| +-- Cumberlandia
+--+
+-- Pseudunio
Lyubas et al., 2019 [Publ 3963], fig. 10
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 3 | 3 | 1 | 3 | 13 | 0 | 0 | 3.0 |
+----- Gibbosula_rochechouartii **
+--+
| | +-- Gibbosula_laosensis **
| +--+
--+ +-- Gibbosula_crassa **
|
| +----- Margaritifera
+--+
| +-- Cumberlandia
+--+
+-- Pseudunio
Campbell & Lydeard, 2026 [Publ 4810], fig. 3_5
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 3 | 3 | 1 | 3 | 137 | 0 | 0 | 3.0 |
+----- Gibbosula_rochechouartii **
+--+
| | +-- Gibbosula_crassa **
| +--+
--+ +-- Gibbosula_laosensis **
|
| +----- Cumberlandia
+--+
| +-- Pseudunio
+--+
+-- Margaritifera
Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 3 | 3 | 1 | 3 | 630 | 0 | 0 | 1.0 |
+-- Cumberlandia
+-----+
| +-- Margaritifera
--+
| +----- Gibbosula_laosensis **
+--+
| +-- Gibbosula_crassa **
+--+
+-- Gibbosula_rochechouartii **
Huang et al., 2018 [Publ 3828], fig. 3_5
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 27 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii **
+-----+
| +-- Gibbosula_laosensis **
--+
| +----- Margaritifera
+--+
| +-- Cumberlandia
+--+
+-- Pseudunio
Huang et al., 2019 [Publ 3843], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 0.000 | 2 | 13 | 0 | 2 | 199 | 0 | 0 | 13.0 |
+-------- Gibbosula_rochechouartii **
|
+--------------------------------+ +----- Gibbosula_laosensis **
| | |
| +--+ +-- Cumberlandia
| | |
| +--+-- Pseudunio
| |
| +-- Margaritifera
|
| +-- Trapezidens
| +-----------------+
| | +-- Lamellidens
| +-----------------+
| | | +----------------- Parreysia
| | | |
| | +--+ +-------------- Leoparreysia
| | | |
| | | | +-------- Indonaia
| | +--+ +--+
| | | | | +----- Indochinella
| | | | +--+
| | | | | +-- Scabies
--+ | +--+ +--+
| | | +-- Radiatula
| | |
| | | +-------- Coelatura_aegyptiaca__2
| | | |
| | +--+ +----- Coelatura_aegyptiaca__1
| | | |
| | | +----- Nitia
| | +--+
| | | +-- Coelatura_gabonensis
| | | |
| | +--+-- Coelatura_kunenensis
| | |
| | +-- Prisodontopsis
| |
| | +-------------------- Acuticosta
| | |
| | | +----------------- Lepidodesma
| | | |
| | | | +-------------- Scriptolamprotula
| | +--------------+ +--+
+--+ | | | | +----------- Unio
| | | | +--+
| | | | | +-------- Schistodesmus
| | | | +--+
| | | | | +----- Nodularia
| | +--+ +--+
| | | | +-- Tchangsinaia
| | | +--+
| | | +-- Cuneopsis
| | |
| | | +-------------- Lanceolaria
| | | |
| | | | +-------- Anemina
| | | | +--+
| | | | | | +----- Cristaria
| | +--+ | +--+
| | | | | +-- Pletholophus
| | | | +--+
| | | | +-- Sinanodonta
| | | |
| | +--+ +-------- Alasmidonta
| | | |
| | | +-------- Lasmigona
| | | |
| | | +-------- Pseudanodonta
| | | |
| | | +-------- Simpsonaias
| | +--+
| | +-------- Strophitus
| | |
| | | +-- Pyganodon
| | +-----+
+--+ | +-- Utterbackia
| |
| | +----- Anodonta_anatina
| +--+
| | +-- Anodonta_cygnea
| +--+
| +-- Brachyanodon
|
| +----- Physunio
| +--+
| | | +-- Yaukthwa
| | +--+
| | +-- Lens
| +-----------------------+
| | | +----- Hyriopsis_desowitzi
| | | |
| | | +----- Rectidens
| | | |
| | +--+----- Ensidens
| | |
| | | +-- Hyriopsis_bialata
| | | |
| | | +-- Hyriopsis_myersiana
| | +--+
| | +-- Hyriopsis_panhai
| | |
| | +-- Hyriopsis_khoratensis
| |
| | +-- Chamberlainia
| | +--------------------+
| | | +-- Sinohyriopsis
| | |
| | | +----- Lamprotula
+--+ | +--------------+
| +-----+ | | +-- Potomida
| | | | +--+
| | | | +-- Pronodularia
| | | |
| | | | +----------- Sinosolenaia
| | | | |
| | +--+ | +----- Gonidea
| | | +-----+ +--+
| | | | | | | +-- Leguminaia
| | | | | | +--+
| | | | +--+ +-- Microcondylaea
| | | | |
| | | | | +-- Ptychorhynchus
| | +--+ | |
| | | +-----+-- Parvasolenaia
| | | |
| | | +-- Koreosolenaia
| | |
| | | +-------------- Indopseudodon
+--+ | |
| +--+ +----------- Pilsbryoconcha
| | |
| +--+ +-- Pseudodon
| | +-----+
| | | +-- Sundadontina
| +--+
| | +----- Nyeinchanconcha
| +--+
| | +-- Thaiconcha
| +--+
| +-- Bineurus
|
| +----- Quadrula
| +--------------------+
| | | +-- Megalonaias
| | +--+
| | +-- Uniomerus
| |
| | +-------- Elliptoideus
+--+ |
| +--------------+ +-- Elliptio
| | | +--+
| | | | +-- Eurynia
| | +--+
| | | +-- Fusconaia
+--+ +--+
| +-- Pleurobema
|
| +-------------------- Amblema
| |
| | +----- Toxolasma
+--+ +-----------+
| | | +-- Obliquaria
| | +--+
| | +-- Plectomerus
+--+
| +-------------- Potamilus_leptodon
| |
| | +----------- Potamilus_alatus
+--+ |
| | +-------- Venustaconcha
| | |
+--+ +-------- Leaunio
| |
| | +-- Ellipsaria
| +-----+
+--+ +-- Truncilla
|
| +-- Medionidus
+-----+
| +-- Ptychobranchus
|
| +----- Cambarunio
+--+
| +-- Ortmanniana
+--+
+-- Lampsilis
Huang et al., 2019 [Publ 3843], fig. S3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 0.000 | 2 | 13 | 0 | 2 | 197 | 0 | 0 | 13.0 |
+----------- Gibbosula_rochechouartii **
+--------------------------------------------+
| | +-------- Gibbosula_laosensis **
| +--+
| | +----- Margaritifera
| +--+
| | +-- Cumberlandia
| +--+
| +-- Pseudunio
|
| +-- Trapezidens
| +--------------------+
| | +-- Lamellidens
| +-----------------------------+
| | | +-------------------- Parreysia
| | | |
| | +--+ +----------------- Leoparreysia
| | | |
--+ | | | +-------- Indonaia
| | +--+ +-----+
| | | | | +----- Indochinella
| | | | +--+
| | | | | +-- Scabies
| | +--+ +--+
| | | +-- Radiatula
| | |
| | | +----------- Coelatura_aegyptiaca__2
| | | |
| | +--+ +-- Coelatura_aegyptiaca__1
| | | +-----+
| | | | +-- Nitia
| | +--+
| | | +----- Coelatura_kunenensis
| | +--+
| | | +-- Coelatura_gabonensis
| | +--+
+--+ +-- Prisodontopsis
|
| +-------------- Indopseudodon
| |
| +-----------------------------------+ +----------- Pilsbryoconcha
| | | |
| | | | +-- Sundadontina_tumida
| | +--+ +-----+
| | | | +-- Pseudodon
| | | |
| | +--+ +-- Nyeinchanconcha
| | | +--+
| | | | +-- Sundadontina_cumingii
| | +--+
| | | +-- Thaiconcha
| | +--+
| | +-- Bineurus
| |
| | +----- Physunio
| | +-----------+
| | | | +-- Yaukthwa
| | | +--+
| | +-----------------------------+ +-- Lens
| | | |
+--+ | | +-------------- Ensidens
| | | |
| | +--+ +----------- Hyriopsis_desowitzi
| | | |
| | +--+ +-------- Rectidens
| | | |
| | +--+ +-- Hyriopsis_panhai
| | | +--+
| | | | +-- Hyriopsis_khoratensis
| | +--+
| | | +-- Hyriopsis_bialata
| | +--+
| | +-- Hyriopsis_myersiana
| |
| | +-------------------------- Acuticosta
| | |
| | | +-------------- Scriptolamprotula
| | | +--------+
| | | | | +----------- Unio
| | +-----------------+ | +--+
| | | | | | +-------- Nodularia
+--+ | | | +--+
| | | | | +----- Schistodesmus
| | | | +--+
| | +--+ | +-- Tchangsinaia
| | | +--+
| | | +-- Cuneopsis
| | |
| | | +-------------------- Lepidodesma
| | | |
| | | | +----------------- Lanceolaria
| | +--+ |
| | | | +-------- Anemina
| | | | +-----+
| | | | | | +----- Cristaria
| | +--+ | +--+
| | | | | +-- Pletholophus
| | | | +--+
| | | | +-- Sinanodonta
| | | |
| | +--+ +-- Alasmidonta
| | | +--------+
| | | | +-- Lasmigona
| | | |
+--+ | | +----- Anodonta_anatina
| | +-----+
| | | | +-- Anodonta_cygnea
| +--+ +--+
| | +-- Brachyanodon
| |
| | +-- Pyganodon
| | +-----+
| | | +-- Utterbackia
| +--+
| | +----- Pseudanodonta
| +--+
| | +-- Simpsonaias
| +--+
| +-- Strophitus
|
| +-- Chamberlainia
| +--------------------------------------+
| | +-- Sinohyriopsis
| |
| | +----- Lamprotula
| | +--------+
| | | | +-- Potomida
| | | +--+
| | | +-- Pronodularia
| | +-----------------------+
+--+ | | +----------- Sinosolenaia
| | | |
| | | | +----- Gonidea
| | +--+ +--+
| | | | | +-- Leguminaia
| | | | +--+
| | +--+ +-- Microcondylaea
| | |
| | | +----- Ptychorhynchus
| | +--+
+--+ | +-- Parvasolenaia
| +--+
| +-- Koreosolenaia
|
| +----- Quadrula
| +-----------------------------+
| | | +-- Megalonaias
| | +--+
| | +-- Uniomerus
| |
| | +-------- Elliptoideus
+--+ |
| +-----------------------+ +-- Elliptio
| | | +--+
| | | | +-- Eurynia
| | +--+
| | | +-- Fusconaia
+--+ +--+
| +-- Pleurobema
|
| +----------------------------- Amblema
| |
+--+ +-------------------------- Toxolasma
| |
+--+ +-- Obliquaria
| +--------------------+
| | +-- Plectomerus
+--+
| +-------------------- Potamilus_leptodon
| |
+--+ +----------------- Potamilus_alatus
| |
| | +-- Medionidus
+--+ +-----------+
| | +-- Ptychobranchus
| |
+--+ +-- Ellipsaria
| +--------+
| | +-- Truncilla
+--+
| +-------- Leaunio
| |
+--+ +-- Venustaconcha
| +--+
| | +-- Cambarunio
+--+
| +-- Ortmanniana
+--+
+-- Lampsilis
Huang et al., 2019 [Publ 3843], fig. S4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 0.000 | 2 | 13 | 0 | 2 | 199 | 0 | 0 | 13.0 |
+----------- Gibbosula_rochechouartii **
+-----------+
| | +-------- Gibbosula_laosensis **
| +--+
| | +----- Cumberlandia
| +--+
| | +-- Pseudunio
| +--+
| +-- Margaritifera
|
| +----------------- Parreysia
| |
| | +-- Trapezidens
| +--------------+
| | +-- Lamellidens
| +--+
| | | +-------------- Leoparreysia
| | | |
| | | | +-------- Indonaia
| | | | +--+
| | +--+ | | +----- Indochinella
| | | | +--+
| | | | | +-- Scabies
| | | | +--+
| | +--+ +-- Radiatula
| | |
| | | +-------- Coelatura_aegyptiaca__2
| | | |
| | | +-------- Nitia
| | +--+
| | | +----- Coelatura_aegyptiaca__1
--+ | | |
| | +--+ +-- Coelatura_gabonensis
| | | |
| | +--+-- Coelatura_kunenensis
| | |
| | +-- Prisodontopsis
| |
| | +----------------- Chamberlainia
| | |
| | +----------------- Megalonaias
| | |
| | +----------------- Uniomerus
| | |
| | +----------------- Quadrula
| | |
| | +----------------- Sinohyriopsis
| | |
| | | +----- Lamprotula
| | +-----------+
| | | | +-- Potomida
| | | +--+
| | | +-- Pronodularia
| | |
| | | +----- Indopseudodon
| | | |
| | | | +-- Nyeinchanconcha
| | | | |
| | +-----------+ +-- Thaiconcha
| | | | |
| | | | +-- Sundadontina_cumingii
+--+ | | |
| | | +-- Sundadontina_tumida__1
| | +--+
| | +-- Sundadontina_tumida__2
| | |
| | +-- Pseudodon
| | |
| | +-- Pilsbryoconcha
| | |
| | +-- Bineurus
| |
| | +-------- Sinosolenaia_carinata
| | |
| | | +----- Gonidea
| | +--+
| | | | +-- Leguminaia
| | | +--+
| | | +-- Microcondylaea
| +--------+
| | | +----- Ptychorhynchus
| | +--+
| | | | +-- Parvasolenaia
| | | +--+
| | | +-- Koreosolenaia
| | |
| | | +----- Sinosolenaia_recognita
| | +--+
| | | +-- Sinosolenaia_iridinea
| | +--+
| | +-- Sinosolenaia_oleivora
| |
| | +-- Physunio_modelli
| | |
| | +-- Physunio
| | |
| | +-----+-- Physunio_superbus
| | | |
| | | +-- Yaukthwa
| | | |
| | | +-- Lens
| +--------+
| | | +----- Hyriopsis_desowitzi
| | | |
| | | +----- Hyriopsis_myersiana
| | | |
| | | +----- Hyriopsis_panhai
| | +--+
+--+ | +-- Hyriopsis_bialata
| +--+
| | +-- Hyriopsis_khoratensis
| |
| | +-- Rectidens
| +--+
| +-- Ensidens
|
| +-------------- Amblema
| |
| +-------------- Toxolasma
| |
| | +-- Obliquaria
| +-----------+
| | +-- Plectomerus
| |
| | +----------- Elliptoideus
| +--+
| | | +-------- Elliptio
| | +--+
| | | +----- Eurynia
+--+ +--+
| | | +-- Fusconaia
| | +--+
| | +-- Pleurobema
| |
| | +----- Ellipsaria
| | |
| | +----- Potamilus_leptodon
| | |
| | +----- Medionidus
| | |
| | +----- Potamilus_alatus
| | |
| | +----- Ptychobranchus
| +--------+
| +----- Truncilla
| |
| +----- Venustaconcha
| |
| +----- Cambarunio
| |
| +----- Leaunio
| |
| | +-- Ortmanniana
| +--+
| +-- Lampsilis
|
| +-------- Acuticosta
| |
| +-------- Alasmidonta
| |
| +-------- Anodonta_anatina
| |
| +-------- Anemina
| |
| +-------- Anodonta_cygnea
| |
| +-------- Cristaria
| |
| +-------- Brachyanodon
| |
| +-------- Lasmigona
| |
| +-------- Lepidodesma
| |
| +-------- Pseudanodonta
| |
| +-------- Pyganodon
| |
+--------+-------- Simpsonaias
|
+-------- Strophitus
|
+-------- Utterbackia
|
+-------- Schistodesmus
|
+-------- Unio
|
+-------- Scriptolamprotula
|
+-------- Lanceolaria
|
| +-- Pletholophus
+-----+
| +-- Sinanodonta
|
| +----- Cuneopsis_celtiformis
| |
| +----- Tchangsinaia
| |
+--+----- Cuneopsis_rufescens
|
+----- Nodularia
|
| +-- Cuneopsis_szechenyii
+--+
+-- Cuneopsis_heudei
Gomes-dos-Santos et al., 2019 [Publ 3943], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 8 | 0 | 0 | -- |
+-- Gibbosula_crassa **
+-----+
| +-- Gibbosula_rochechouartii **
--+
| +----- Cumberlandia
+--+
| +-- Pseudunio
+--+
+-- Margaritifera
Wu et al., 2020 [Publ 4121], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 32 | 0 | 0 | -- |
+----- Pseudunio --+ | +-- Gibbosula_laosensis ** +--+ +-- Gibbosula_rochechouartii **
Wu et al., 2020 [Publ 4121], fig. S1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 0.000 | 2 | 12 | 0 | 2 | 32 | 0 | 0 | 12.0 |
+----------- Gibbosula_rochechouartii **
+--+
| | +-------- Gibbosula_laosensis **
| +--+
| | +----- Margaritifera
| +--+
| | +-- Cumberlandia
--+ +--+
| +-- Pseudunio
|
| +-- Lampsilis
| +--------+
| | +-- Scabies
| |
+--+ +-- Unio
| +-----+
| | +-- Scriptolamprotula
+--+
| +----- Sinosolenaia
+--+
| +-- Potomida
+--+
+-- Lamprotula
Sano et al., 2020 [Publ 4138], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 67 | 0 | 0 | -- |
+----- Pseudunio --+ | +-- Gibbosula_rochechouartii ** +--+ +-- Gibbosula_laosensis **
Sano et al., 2020 [Publ 4138], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 67 | 0 | 0 | -- |
+----- Margaritifera --+ | +-- Gibbosula_laosensis ** +--+ +-- Gibbosula_rochechouartii **
Sano et al., 2020 [Publ 4138], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 67 | 0 | 0 | -- |
+----- Margaritifera --+ | +-- Gibbosula_rochechouartii ** +--+ +-- Gibbosula_laosensis **
Wu et al., 2021 [Publ 4366], fig. 3-BI
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 71 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii **
+-----+
| +-- Gibbosula_crassa **
--+
| +----- Cumberlandia
+--+
| +-- Pseudunio
+--+
+-- Margaritifera
Wu et al., 2021 [Publ 4366], fig. 3-ML
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 71 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii **
+-----+
| +-- Gibbosula_crassa **
--+
| +----- Cumberlandia
+--+
| +-- Pseudunio
+--+
+-- Margaritifera
Wu et al., 2021 [Publ 4366], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 66 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii **
+-----+
| +-- Gibbosula_crassa **
--+
| +----- Cumberlandia
+--+
| +-- Pseudunio
+--+
+-- Margaritifera
Wu et al., 2023 [Publ 4339], fig. 4_S5-aa
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 93 | 0 | 0 | -- |
+-- Gibbosula_crassa **
+-----+
| +-- Gibbosula_rochechouartii **
--+
| +----- Cumberlandia
+--+
| +-- Pseudunio
+--+
+-- Margaritifera
Wu et al., 2023 [Publ 4339], fig. 4_S5-nt
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 93 | 0 | 0 | -- |
+-- Gibbosula_crassa **
+-----+
| +-- Gibbosula_rochechouartii **
--+
| +----- Cumberlandia
+--+
| +-- Pseudunio
+--+
+-- Margaritifera
Chen et al., 2026 [Publ 4833], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 2 | 2 | 0 | 2 | 25 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii **
+--------------------------------+
| +-- Gibbosula_sinensis **
--+
| +-------------------------------- Lamellidens
| |
+--+ +----------------------------- Trapezidens
| |
+--+ +-------------------------- Parreysia
| |
+--+ +----------------------- Leoparreysia
| |
| | +----- Nitia
+--+ +--------------+
| | | +-- Coelatura
| | +--+
| | +-- Prisodontopsis
+--+
| +-- Indochinella
| +--------------+
| | +-- Indonaia
+--+
| +-------------- Radiatula
| |
+--+ +----------- Songkhlanaia
| |
+--+ +-------- Harmandia
| |
+--+ +-- Unionetta_xishuangbannaensis
| +--+
| | +-- Unionetta_fabagina__2
+--+
| +-- Unionetta_fabagina__1
+--+
+-- Scabies
Zhou et al., 2007 [Publ 2658], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 20 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Lamprotula
Bolotov et al., 2016 [Publ 3732], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 7 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Bolotov et al., 2016 [Publ 3743], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 29 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Cumberlandia
Bolotov et al., 2016 [Publ 3743], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 29 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Cumberlandia
Araujo et al., 2017 [Publ 3746], fig. 1_2b
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 12 | 0 | 0 | -- |
+-------- Gibbosula_laosensis ** --+ | +----- Margaritifera +--+ | +-- Cumberlandia +--+ +-- Pseudunio
Araujo et al., 2017 [Publ 3746], fig. 2a
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 10 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Bolotov et al., 2017 [Publ 3761], fig. S1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 138 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Pseudunio
Bolotov et al., 2017 [Publ 3761], fig. S2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 138 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** | --+-- Pseudunio | +-- Margaritifera
Vikhrev et al., 2017 [Publ 3810], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 9 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Bespalaya et al., 2018 [Publ 3811], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 9 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Vikhrev et al., 2018 [Publ 3812], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 12 | 0 | 0 | -- |
+-------- Gibbosula_laosensis ** --+ | +----- Margaritifera +--+ | +-- Cumberlandia +--+ +-- Pseudunio
Huang et al., 2018 [Publ 3828], fig. 6
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 41 | 0 | 0 | -- |
+-------- Gibbosula_rochechouartii ** --+ | +----- Cumberlandia +--+ | +-- Pseudunio +--+ +-- Margaritifera
Huang et al., 2019 [Publ 3843], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 45 | 0 | 0 | -- |
+-- Margaritifera --+ +-- Gibbosula_rochechouartii **
Huang et al., 2019 [Publ 3843], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 55 | 0 | 0 | -- |
+-------- Gibbosula_rochechouartii ** --+ | +----- Cumberlandia +--+ | +-- Pseudunio +--+ +-- Margaritifera
Huang et al., 2019 [Publ 3843], fig. S1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 46 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Wu et al., 2019 [Publ 3864], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 45 | 0 | 0 | -- |
+----- Gibbosula_rochechouartii ** --+ | +-- Cumberlandia +--+ +-- Margaritifera
Konopleva et al., 2019 [Publ 3900], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 50 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Guerra et al., 2019 [Publ 4168], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 40 | 0 | 0 | -- |
+-------- Gibbosula_crassa ** --+ | +----- Cumberlandia +--+ | +-- Pseudunio +--+ +-- Margaritifera
Guerra et al., 2019 [Publ 4168], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 40 | 0 | 0 | -- |
+-------- Gibbosula_crassa ** --+ | +----- Cumberlandia +--+ | +-- Pseudunio +--+ +-- Margaritifera
Bolotov et al., 2020 [Publ 4100], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 116 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** | +-- Margaritifera_dahurica | --+-- Margaritifera_margaritifera | +-- Margaritifera_laevis | +-- Margaritifera_middendorffi
Fukata & Iigo, 2020 [Publ 4106], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 53 | 0 | 0 | -- |
+----- Gibbosula_crassa ** --+ | +-- Pseudunio +--+ +-- Cumberlandia
Wu et al., 2020 [Publ 4121], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 47 | 0 | 0 | -- |
+----- Gibbosula_rochechouartii ** --+ | +-- Cumberlandia +--+ +-- Margaritifera
Sano et al., 2020 [Publ 4138], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 33 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Wu et al., 2020 [Publ 4147], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 13 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Chen et al., 2020 [Publ 4170], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 33 | 0 | 0 | -- |
+----------------- Gibbosula_crassa ** --+ | +-------------- Quadrula +--+ | +----------- Toxolasma +--+ | +-------- Lampsilis_ornata +--+ | +----- Potamilus +--+ | +-- Lampsilis_siliquoidea +--+ +-- Lampsilis_powellii
Wu et al., 2021 [Publ 4366], fig. 2_I
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 90 | 0 | 0 | -- |
+-------- Gibbosula_laosensis ** --+ | +----- Margaritifera +--+ | +-- Pseudunio +--+ +-- Cumberlandia
Wu et al., 2021 [Publ 4366], fig. 6
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 90 | 0 | 0 | -- |
+-------- Gibbosula_laosensis ** --+ | +----- Margaritifera +--+ | +-- Cumberlandia +--+ +-- Pseudunio
Wu et al., 2022 [Publ 4238], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 72 | 0 | 0 | -- |
+-------- Gibbosula_rochechouartii ** --+ | +----- Cumberlandia +--+ | +-- Pseudunio +--+ +-- Margaritifera
Bolotov et al., 2022 [Publ 4250], fig. 8
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 48 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** | +-- Margaritifera | +-- Sinosolenaia | --+-- Leguminaia | +-- Gonidea | +-- Lamprotula | +-- Potomida
Jeratthitikul et al., 2022 [Publ 4260], fig. S1_1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 75 | 0 | 0 | -- |
+----- Gibbosula_laosensis ** --+ | +-- Pseudunio +--+ +-- Margaritifera
Jeratthitikul et al., 2022 [Publ 4260], fig. S2_2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 75 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Pseudunio
Konopleva et al., 2023 [Publ 4269], fig. S1_4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 55 | 0 | 0 | -- |
+----------------------------- Gibbosula_laosensis ** | | +-- Trapezidens | +-----------+ | | +-- Lamellidens | +-----------+ | | | +----------- Parreysia | | +--+ | | | +-------- Leoparreysia | | +--+ --+ | | +----- Indonaia | | +--+ | | | +-- Indochinella | | +--+ | | +-- Radiatula | | | | +-- Lampsilis | | +--------------+ | | | +-- Amblema | | | +--+ | +-- Chamberlainia | | +-----+ | +-----+ | +-- Sinohyriopsis | | | +-----+ | | | | | +----- Potomida | | | | +--+ | | | | | +-- Pronodularia | | +--+ +--+ | | | +-- Lamprotula | | | | | | +----------- Sinosolenaia | | | | | | +--+ +-- Ptychorhynchus | | | +-----+ | | | | +-- Parvasolenaia | | +--+ +--+ | +----- Gonidea | +--+ | | +-- Microcondylaea | +--+ | +-- Leguminaia | | +-------------- Middendorffinaia | +-----+ | | | +----------- Scriptolamprotula | | +--+ | | | +-------- Unio | | +--+ | | | +----- Nodularia | | +--+ +--+ | +-- Tchangsinaia | +--+ | +-- Cuneopsis | | +-- Lepidodesma | +--------------+ | | +-- Lanceolaria | | +--+ +-------------- Acuticosta | | | | +----- Anodonta_anatina +--+ +-----+ | | | +-- Pseudanodonta | | +--+ | | +-- Anodonta_cygnea +--+ | +----- Anemina | +--+ | | | +-- Amuranodonta | | +--+ +--+ +-- Buldowskia | | +----- Cristaria +--+ | +-- Beringiana +--+ +-- Sinanodonta
Dai et al., 2023 [Publ 4285], fig. 3-BI
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 44 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Dai et al., 2023 [Publ 4285], fig. 3-ML
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 44 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Dai et al., 2023 [Publ 4285], fig. 5
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 52 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Bogan et al., 2023 [Publ 4295], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 30 | 0 | 0 | -- |
+-- Gibbosula_crassa ** --+ +-- Margaritifera
Bogan et al., 2023 [Publ 4295], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 6 | 0 | 0 | -- |
+-------- Gibbosula_crassa ** --+ | +----- Cristaria +--+ | +-- Sinanodonta +--+ +-- Pletholophus
Bogan et al., 2023 [Publ 4295], fig. 5
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 5 | 0 | 0 | -- |
+-------- Gibbosula_crassa ** --+ | +----- Cristaria +--+ | +-- Sinanodonta +--+ +-- Pletholophus
Bogan et al., 2023 [Publ 4295], fig. 6
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 12 | 0 | 0 | -- |
+-------- Gibbosula_crassa ** --+ | +----- Pletholophus +--+ | +-- Cristaria +--+ +-- Sinanodonta
Chen et al., 2023 [Publ 4302], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 40 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Chen et al., 2023 [Publ 4302], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 40 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Dai et al., 2024 [Publ 4358], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 52 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Wu et al., 2024 [Publ 4379], fig. 5
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 52 | 0 | 0 | -- |
+-- Gibbosula_crassa ** --+ +-- Margaritifera
Dai et al., 2024 [Publ 4385], fig. 3-BI
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 30 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Dai et al., 2024 [Publ 4385], fig. 3-ML
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 30 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Dai et al., 2024 [Publ 4386], fig. 5-BI
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 21 | 0 | 0 | -- |
+-- Margaritifera --+ +-- Gibbosula_laosensis **
Dai et al., 2024 [Publ 4386], fig. 5-ML
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 21 | 0 | 0 | -- |
+-- Margaritifera --+ +-- Gibbosula_laosensis **
Dai et al., 2024 [Publ 4389], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 45 | 0 | 0 | -- |
+-- Margaritifera --+ +-- Gibbosula_laosensis **
Kashiri et al., 2024 [Publ 4822], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 9 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Dai et al., 2025 [Publ 4440], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 43 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Dai et al., 2025 [Publ 4440], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 43 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Dai et al., 2025 [Publ 4440], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 86 | 0 | 0 | -- |
+-------- Gibbosula_rochechouartii ** --+ | +----- Cumberlandia +--+ | +-- Pseudunio +--+ +-- Margaritifera
Dai et al., 2025 [Publ 4441], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 46 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Chen et al., 2025 [Publ 4448], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 84 | 0 | 0 | -- |
+-------- Gibbosula_rochechouartii ** --+ | +----- Cumberlandia +--+ | +-- Pseudunio +--+ +-- Margaritifera
Chen et al., 2025 [Publ 4487], fig. 2
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 40 | 0 | 0 | -- |
+-- Margaritifera --+ +-- Gibbosula_rochechouartii **
Chen et al., 2025 [Publ 4487], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 40 | 0 | 0 | -- |
+-- Margaritifera --+ +-- Gibbosula_rochechouartii **
Wang et al., 2025 [Publ 4488], fig. 3B
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 63 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Chen et al., 2025 [Publ 4663], fig. 1
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 47 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Dai et al., 2026 [Publ 4757], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 33 | 0 | 0 | -- |
+-- Gibbosula_laosensis ** --+ +-- Margaritifera
Wu et al., 2026 [Publ 4811], fig. 3
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 16 | 0 | 0 | -- |
+-- Margaritifera --+ +-- Gibbosula_rochechouartii **
Wu et al., 2026 [Publ 4811], fig. 4
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 94 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
Wu et al., 2026 [Publ 4825], fig. 6B
| iT | tT | bi | riS | oT | roT | roS | iD | |
| 1.000 | 1 | 1 | 0 | 1 | 64 | 0 | 0 | -- |
+-- Gibbosula_rochechouartii ** --+ +-- Margaritifera
