The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)
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Bivalvia | Unionida | Unionoidea | Unionidae | Ambleminae tribe QUADRULINI Ihering, 1901: cladomics
5 genera, 1 subtribe Cladomics Summarynumber of species = 26 (46 total)
number of subtaxa (rank: subtribe) = 6
analyzed in 201 cladograms in 121 publications
maximum number of terminals in an analysis =
22 (mean = 2.64, median = 1)
maximum number of subtaxa (subtribe) =
3 (mean = 1.36, median = 1) Basic Monophylynumber of cladograms that test basic monophyly
(i.e., more than 1 included species) = 89
maximum number of included terminals = 22, (mean =
4.71 median = 2)
maximum number of subtaxa (subtribe) =
3 (mean = 1.82, median = 2)
proportion of cladograms that support basic monophyly
= 0.83 Rigorous Monophylynumber of cladograms that test rigorous monophyly = 52
(Rigorous monophyly refers to cladograms with terminal species representing
more than one subtaxon (subtribe) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 22 (mean = 5.96, median = 4)
maximum number of subtaxa (subtribe) =
3 (mean = 2.35, median = 2)
proportion of cladograms that support rigorous monophyly = 0.77
Pairwise Cladogram ClusteringThe graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.
Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14 ** Top Monophyly Cladogram ** ** Top Sister-Group Cladogram ** iT| tT | bi | riS | oT | roT | roS | iD | | 21 | 21 | 21 | 3 | 611 | 170 | 4 | 0 | +-------- Pleurobemini
+--------------------------------------------------------------------+
| | +----- Quadrulina **
| +--+
| | +-- Megalonaias **
| +--+
| +-- Uniomerus **
--+
| +----- Amblema
| +--------------------------------------------------------------------+
| | | +-- Nephronaias
| | +--+
| | +-- Psoronaias
| |
+--+ +----------------------------------------------------------------------- Reginaia
| |
| | +----- Popenaias
| | +--------------------------------------------------------------+
| | | | +-- Cyrtonaias
+--+ | +--+
| | +-- Glebula
| |
| | +----------------------------------------------------------------- Paetulunio
| | |
+--+ | +----------------- Toxolasma_parvum
| | |
| | | +----- Toxolasma_lividum__2
| | | +--------+
| | +--------------------------------------------+ | | +-- Toxolasma_cylindrellus
| | | | | +--+
| | | | | +-- Toxolasma_lividum__1
| | | | |
+--+ | +--+ +-------- Toxolasma_texasiense__2
| | | +--+
| | | | | +----- Toxolasma_glans
| | | | +--+
| | | | | +-- Toxolasma_paulum
| | +--+ +--+
| | | +-- Toxolasma_texasiense__1
| | |
| | | +-------- Toxolasma__4
| | +--+
+--+ | +----- Toxolasma_granulatum
| +--+
| | +-- Toxolasma__2
| +--+
| +-- Toxolasma__3
|
| +----- Toxolasma_pullus
| +-----------------------------------------------------+
| | | +-- Toxolasma_germanum
| | +--+
| | +-- Toxolasma__1
| |
| | +-- Obliquaria
| | +-----------------------------------------------------+
| | | +-- Plectomerus
+--+ |
| | +-------- Potamilus_inflatus
| | +--------+
| | | | +----- Potamilus_ohiensis
| | | +--+
| | | | +-- Potamilus_amphichaenus
| | | +--+
| | +-----------------------------------+ +-- Potamilus_streckersoni
| | | |
+--+ | | +----- Potamilus_fragilis__2
| | | +--------+
| | | | | +-- Potamilus_leptodon
| | | | +--+
| | +--+ +-- Potamilus_fragilis__1
| | |
| | | +----------- Potamilus_metnecktayi
| | | |
| | +--+ +-------- Potamilus_alatus__3
| | | |
| | +--+ +----- Potamilus_alatus__2
+--+ | |
| +--+----- Potamilus_purpuratus__2
| |
| | +-- Potamilus_alatus__1
| +--+
| +-- Potamilus_purpuratus__1
|
| +-- Villosa_villosa
| +-----------------------------------------------+
| | +-- Atlanticoncha
| |
| | +-- Cambarunio_nebulosus
| | +--------------------------------------------+
+--+ | +-- Ligumia
| |
| | +-------- Leaunio_umbrans
| | +-----------------------------------+
| | | | +----- Cambarunio_taeniatus
+--+ | +--+
| | | +-- Cambarunio_iris
| | +--+
| | +-- Cambarunio_simus
| |
| | +----------------------------------------- Hamiota
| | |
+--+ | +----------------------- Pachynaias
| | |
| | +--------------+ +-------------------- Sagittunio
| | | | |
| | | | | +----- Ellipsaria
| | | +--+ +-----------+
| | | | | | +-- Potamilus_capax
| | | | | +--+
| | | | | +-- Truncilla
+--+ | +--+
| | | +----- Medionidus_acutissimus
| | | +--------+
| | | | | +-- Lemiox
| | | | +--+
| | +--+ +-- Medionidus_penicillatus
| | |
| | | +----------- Ptychobranchus
| | | |
| | +--+ +-- Cyprogenia_aberti__2
| | | +-----+
+--+ | | +-- Dromus
| +--+
| | +----- Cyprogenia_stegaria__2
| +--+
| | +-- Cyprogenia_aberti__1
| +--+
| +-- Cyprogenia_stegaria__1
|
| +-- Villosa_delumbis
| +--------------------------------+
| | +-- Villosa_vibex
| |
| | +-------------- Obovaria_olivaria
| | |
| | +-----------------+ +-- Venustaconcha
| | | | +--------+
| | | | | +-- Epioblasma
| | | +--+
| | | | +-------- Obovaria_retusa
+--+ | +--+
| | | +----- Obovaria_choctawensis
| | +--+
| | | +-- Obovaria_unicolor
| | +--+
| | +-- Obovaria_subrotunda
| |
| | +-- Mesonaias
| | +-----------------------+
| | | +-- Obovaria_arkansasensis
| | |
+--+ +--+ +-- Lampsilis_splendida
| | | +--------------------+
| | | | +-- Lampsilis_bracteata
| | | |
| | +--+ +-- Lampsilis_fasciola
| | | +-----------------+
| | | | +-- Ortmanniana_pectorosa
| | +--+
| | | +----------------- Lampsilis_ornata
| | | |
| | +--+ +-------------- Lampsilis_ovata__2
| | | |
| | +--+ +----------- Lampsilis_dolabraeformis
| | | |
| | | | +----- Lampsilis_cardium__2
| | +--+ +--+
| | | | | +-- Lampsilis_cardium__1
+--+ | | +--+
| +--+ +-- Lampsilis_satura
| |
| | +----- Lampsilis_cariosa__2
| +--+
| | +-- Lampsilis_ovata__1
| +--+
| +-- Lampsilis_cariosa__1
|
| +----------- Ortmanniana_ligamentina__2
| +--------------+
| | | +-------- Lampsilis_abrupta
| | +--+
| | | +----- Lampsilis_higginsii__2
| | +--+
| | | +-- Ortmanniana_ligamentina__1
| | +--+
+--+ +-- Lampsilis_higginsii__1
|
| +----- Medionidus_conradicus
| +-----------------+
| | | +-- Leaunio_lienosus
| | +--+
| | +-- Leaunio_vanuxemensis
| |
+--+ +-- Lampsilis_floridensis
| +-----------------+
| | +-- Lampsilis_teres
| |
| | +-- Lampsilis_siliquoidea__1
+--+ |
| +-- Lampsilis_siliquoidea__2
| +--------------+
| | +-- Lampsilis_powellii__1
| | |
+--+ +-- Lampsilis_powellii__2
|
| +-------------- Lampsilis_sietmani
| |
+--+ +-- Lampsilis_straminea__2
| +--------+
| | +-- Lampsilis_virescens
+--+
| +-------- Lampsilis_hydiana__2
+--+
| +----- Lampsilis_hydiana__1
+--+
| +-- Lampsilis_bergmanni
+--+
+-- Lampsilis_straminea__1
Neemuchwala et al., 2023 [Publ 4292], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 22 | 22 | 20 | 3 | 5 | 4 | 4 | 25 | +----- Megalonaias **
+-----+
| | +-- Uniomerus **
| +--+
--+ +-- Quadrulina **
|
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Lampsilini
+--+
+-- Popenaiadini
Lopes-Lima et al., 2019 [Publ 3901], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 19 | 25 | 23 | 3 | 10 | 6 | 3 | 26 | +----------- Megalonaias **
|
--+ +-- Uniomerus **
| +-----+
| | +-- Quadrulina **
+--+
| +----- Pleurobemini
+--+
| +-- Lampsilini
+--+
+-- Amblemini
Lopes-Lima et al., 2019 [Publ 3901], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 18 | 18 | 15 | 3 | 10 | 6 | 3 | 10 | +-------------- Amblemini
--+
| +----------- Lampsilini
+--+
| +-------- Pleurobemini
+--+
| +----- Quadrulina **
+--+
| +-- Megalonaias **
+--+
+-- Uniomerus **
Lopes-Lima et al., 2019 [Publ 3901], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 16 | 16 | 14 | 3 | 10 | 6 | 3 | 17 | +----- Amblemini
+--+
| | +-- Pleurobemini
| +--+
--+ +-- Lampsilini
|
| +----- Megalonaias **
+--+
| +-- Uniomerus **
+--+
+-- Quadrulina **
Serb et al., 2003 [Publ 2659], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 14 | 14 | 12 | 2 | 15 | 15 | 3 | 21 | +-------------------------- Lampsilis
|
--+ +----------------------- Potamilus
| |
| | +----- Medionidus
+--+ +--------------+
| | | +-- Dromus
| | +--+
+--+ +-- Cyprogenia
|
| +----------------- Obliquaria
| |
+--+ +-------------- Amblema
| |
| | +----- Reginaia
+--+ +-----+
| | | +-- Elliptoideus
| | +--+
| | +-- Plectomerus
+--+
| +-- Megalonaias **
| +-----+
| | +-- Quadrulina **
+--+
| +----- Hemistena
+--+
| +-- Elliptio
+--+
+-- Fusconaia
Johnson et al., 2018 [Publ 3844], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 14 | 14 | 12 | 3 | 2 | 2 | 2 | 17 | +----------- Amblemini
--+
| +-------- Pleurobemini
+--+
| +----- Megalonaias **
+--+
| +-- Uniomerus **
+--+
+-- Quadrulina **
Szumowski et al., 2012 [Publ 3569], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 14 | 14 | 12 | 2 | 0 | 0 | 0 | 25 | +----- Megalonaias **
|
--+----- Theliderma **
|
| +-- Quadrula **
+--+
+-- Pustulosa **
Serb et al., 2003 [Publ 2659], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 14 | 14 | 11 | 2 | 15 | 15 | 3 | 22 | +----------- Lampsilis
|
| +-------- Obliquaria
| |
| +-------- Amblema
| |
--+ +-------- Reginaia
| |
| +-------- Potamilus
| |
| | +-- Elliptoideus
| +-----+
| | +-- Plectomerus
+--+
| +-- Megalonaias **
+-----+
| +-- Quadrulina **
|
| +----- Medionidus
+--+
| | +-- Dromus
| +--+
| +-- Cyprogenia
|
| +----- Hemistena
+--+
| +-- Elliptio
+--+
+-- Fusconaia
Campbell & Lydeard, 2012 [Publ 3309], fig. 1-2 iT| tT | bi | riS | oT | roT | roS | iD | | 11 | 11 | 9 | 2 | 62 | 62 | 4 | 18 | +-- Uniomerus **
+-----------+
| +-- Quadrulina **
--+
| +----------- Pleurobemini
| |
+--+ +-- Plectomerus
| +-----+
| | +-- Reginaia
+--+
| +----- Lampsilini
+--+
| +-- Popenaiadini
+--+
+-- Amblema
Campbell & Lydeard, 2012 [Publ 3309], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 11 | 11 | 8 | 2 | 62 | 62 | 4 | 15 | +-- Uniomerus **
+--------------------------------+
| +-- Quadrulina **
|
--+ +-------------------------------- Pleurobemini
| |
| | +-- Amblema
+--+ +--------------------------+
| | +-- Reginaia
| |
+--+ +-------------------------- Popenaiadini
| |
| | +-- Glebula
+--+ +--------------------+
| | +-- Paetulunio
| |
+--+ +-- Toxolasma
| +-----------------+
| | +-- Plectomerus
| |
+--+ +----------------- Obliquaria
| |
| | +----- Potamilus
+--+ +--------+
| | | +-- Dromus
| | +--+
+--+ +-- Lemiox
|
| +----------- Lampsilis_ovata
| |
+--+ +-------- Lampsilis_virescens
| |
+--+ +-- Epioblasma
| +--+
| | +-- Obovaria_arkansasensis
+--+
| +-- Obovaria_olivaria
+--+
+-- Venustaconcha
Campbell et al., 2005 [Publ 2660], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 11 | 11 | 8 | 3 | 91 | 79 | 4 | 17 | +----- Quadrulina **
+--------+
| | +-- Megalonaias **
| +--+
--+ +-- Uniomerus **
|
| +----------- Pleurobemini
| |
+--+ +-- Plectomerus
| +-----+
| | +-- Reginaia
+--+
| +----- Lampsilini
+--+
| +-- Popenaiadini
+--+
+-- Amblema
Campbell et al., 2005 [Publ 2660], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 11 | 11 | 8 | 3 | 91 | 79 | 4 | 19 | +----- Megalonaias **
+--------------------+
| | +-- Uniomerus **
| +--+
| +-- Quadrulina **
|
--+ +----------------------- Pleurobemini
| |
| | +----- Popenaiadini
| | +--------------+
| | | | +-- Amblema
+--+ | +--+
| | +-- Reginaia
| |
| | +-- Plectomerus
| | +--------------+
+--+ | +-- Toxolasma
| |
| | +----- Obliquaria
| | +--------+
| | | | +-- Cyrtonaias
| | | +--+
| | | +-- Glebula
| | |
+--+ | +----------- Sagittunio
| | |
| | +----------- Leaunio
| | |
| | +----------- Potamilus
| | |
| | +----------- Hamiota
| | |
| | | +-- Ellipsaria
+--+ +--------+
| | +-- Truncilla
| |
| | +----- Lampsilis_teres
| +-----+
| | | +-- Ortmanniana_ligamentina
| | +--+
| | +-- Lampsilis_siliquoidea
| |
| | +----- Villosa
| +-----+
| | | +-- Ligumia
| | +--+
| | +-- Cambarunio
| |
+--+ +-------- Ortmanniana_pectorosa
+--+
| | +----- Lampsilis_ornata
| +--+
| | +-- Lampsilis_cardium
| +--+
| +-- Lampsilis_ovata
|
| +-------- Obovaria_olivaria
| |
| +-------- Venustaconcha_ellipsiformis__2
+--+
| +-------- Epioblasma
| |
| | +----- Obovaria_unicolor
| +--+
| | +-- Obovaria_arkansasensis
| +--+
| +-- Obovaria_subrotunda
|
| +-- Ptychobranchus
| +-----+
| | +-- Venustaconcha_ellipsiformis__1
| |
+--+ +----- Medionidus_conradicus
| |
| | +-- Lemiox
+--+--+
| +-- Medionidus_acutissimus
|
| +-- Dromus
+--+
+-- Cyprogenia
Inoue et al., 2018 [Publ 3858], fig. 1_3a-c iT| tT | bi | riS | oT | roT | roS | iD | | 7 | 7 | 2 | 2 | 98 | 76 | 4 | 20 | +-------- Reginaia
|
+-------- Amblema
|
| +----- Pleurobemini
+--+
| | +-- Uniomerus **
| +--+
--+ +-- Quadrulina **
|
| +----- Popenaiadini
| |
| | +-- Dromus
| | |
| | +-- Epioblasma
| | |
+--+ +-- Glebula
| |
| +-- Lampsilis_ovata
| |
| +-- Lampsilis_virescens
| |
| +-- Lemiox
+--+
+-- Obliquaria
|
+-- Obovaria_olivaria
|
+-- Obovaria_unicolor
|
+-- Plectomerus
|
+-- Potamilus_alatus
|
+-- Potamilus_capax
Campbell & Lydeard, 2012 [Publ 3308], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 7 | 7 | 5 | 2 | 36 | 36 | 4 | 18 | +-- Uniomerus **
+-----------------------------+
| +-- Quadrulina **
|
| +-------------- Elliptoideus
| +--------------+
| | | +----------- Plectomerus
--+ | +--+
| | | +-------- Obovaria_unicolor
| | +--+
| | | +----- Obovaria_olivaria
| | +--+
| | | +-- Lampsilis
+--+ +--+
| +-- Lemiox
|
| +-------- Cyrtonaias
| +-----------------+
| | | +----- Reginaia
| | +--+
| | | +-- Popenaiadini
+--+ +--+
| +-- Amblema
|
| +----------------------- Elliptio
| |
+--+ +-------------------- Eurynia
| |
| | +----------------- Parvaspina
+--+ |
| | +-- Hemistena
| +--------------+
+--+ +-- Plethobasus
|
| +-------------- Pleuronaia
| |
| | +-- Pleurobema_cordatum
+--+ +--------+
| | +-- Fusconaia
| |
+--+ +-- Pleurobema_stabile
| +-----+
| | +-- Pleurobema_sintoxia
+--+
| +----- Pleurobema_decisum
+--+
| +-- Pleurobema_clava
+--+
+-- Pleurobema_pyriforme
Zhou et al., 2007 [Publ 2658], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 7 | 7 | 5 | 1 | 11 | 0 | 0 | 27 | Boyer et al., 2011 [Publ 2718], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 7 | 7 | 5 | 2 | 28 | 22 | 3 | 22 | +----------- Reginaia
|
--+ +-- Amblema
| +-----+
| | +-- Lampsilini
+--+
| +----- Pleurobemini
+--+
| +-- Megalonaias **
+--+
+-- Quadrulina **
Boyer et al., 2011 [Publ 2718], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 7 | 29 | 25 | 2 | 31 | 22 | 3 | 40 | +-------------------------------- Quadrulina **
|
--+ +----------------------------- Megalonaias **
| |
+--+ +-------------------------- Toxolasma
| |
+--+ +----------------------- Amblema
| |
| | +-------------------- Pleurobemini
+--+ |
| | +-- Reginaia
| +-----------------+
+--+ +-- Obliquaria
|
| +----------------- Potamilus
| |
| | +-- Ellipsaria
+--+ +-----------+
| | +-- Truncilla
| |
+--+ +----------- Lampsilis_cardium
| |
| | +----- Ligumia
| +-----+
+--+ | +-- Obovaria
| +--+
| +-- Epioblasma
|
| +-------- Lampsilis_teres
+--+
| +----- Lampsilis_siliquoidea
+--+
| +-- Ortmanniana
+--+
+-- Lampsilis_higginsii
Campbell & Lydeard, 2012 [Publ 3308], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 6 | 6 | 4 | 1 | 36 | 36 | 4 | 17 | +-------------------- Quadrulina **
|
--+ +----------------- Pleurobemini
| |
| | +-- Popenaiadini
+--+ +-----------+
| | +-- Amblema
| |
+--+ +-- Cyrtonaias
| +--------+
| | +-- Reginaia
+--+
| +-------- Plectomerus
+--+
| +----- Lemiox
+--+
| +-- Lampsilis
+--+
+-- Obovaria
Campbell & Lydeard, 2012 [Publ 3308], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 6 | 6 | 4 | 1 | 36 | 36 | 4 | 17 | +-------------------- Quadrulina **
|
--+ +----------------- Pleurobemini
| |
+--+ +-- Reginaia
| +-----------+
| | +-- Amblema
+--+
| +----------- Popenaiadini
| |
+--+ +-------- Cyrtonaias
| |
+--+-------- Plectomerus
|
| +----- Lemiox
+--+
| +-- Lampsilis
+--+
+-- Obovaria
Campbell & Lydeard, 2012 [Publ 3308], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 6 | 6 | 4 | 1 | 36 | 36 | 4 | 17 | +----------------- Quadrulina **
|
--+ +-------------- Pleurobemini
| |
| | +-- Popenaiadini
+--+ +--------+
| | +-- Amblema
| |
+--+ +-------- Plectomerus
| |
| | +-- Cyrtonaias
+--+-----+
| +-- Reginaia
|
| +----- Lemiox
+--+
| +-- Lampsilis
+--+
+-- Obovaria
Pfeiffer et al., 2016 [Publ 3660], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 6 | 6 | 4 | 2 | 32 | 31 | 3 | 21 | +----------------------------- Amblemini
|
--+ +-------------------------- Lampsilini
| |
+--+ +----------------------- Elliptoideus
| |
| | +-- Uniomerus **
+--+ +-----------------+
| | +-- Quadrulina **
| |
+--+ +----------------- Elliptio
| |
| | +-------------- Parvaspina
+--+ |
| +-------------- Plethobasus
| |
| | +-- Hemistena
+--+-----------+
| +-- Eurynia
|
| +----------- Fusconaia
| |
| | +----- Pleurobema_riddellii
+--+ +--+
| | | +-- Pleurobema_sintoxia
| | +--+
+--+ +-- Pleurobema_cordatum
|
| +----- Pleuronaia
| |
+--+ +-- Pleurobema_pyriforme
| |
+--+-- Pleurobema_strodeanum
|
+-- Pleurobema_clava
Pfeiffer et al., 2019 [Publ 3866], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 5 | 5 | 3 | 3 | 26 | 25 | 4 | 22 | +----- Megalonaias **
+-----+
| | +-- Uniomerus **
| +--+
--+ +-- Quadrulina **
|
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Breton et al., 2011 [Publ 3376], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 33 | 32 | 2 | 34 | 15 | 3 | 47 | +-------------- Quadrulina **
|
| +----- Lamprotulini
--+ +-----+
| | | +-- Chamberlainiini
| | +--+
+--+ +-- Anodontini
|
| +-------- Megalonaias **
+--+
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Rosenberg et al., 1997 [Publ 2720], fig. 2A iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 17 | 0 | 3 | 18 | 8 | 3 | 33 | +-- Gonideini
|
+-- Pyganodon_grandis
|
+-- Utterbackia
|
+-- Pyganodon_cataracta
|
+-- Lampsilis_straminea
|
+-- Lampsilis_teres
|
+-- Megalonaias **
|
+-- Obliquaria
|
--+-- Theliderma **
|
+-- Quadrula **
|
+-- Uniomerus **
|
+-- Amblema
|
+-- Elliptio
|
+-- Fusconaia
|
+-- Plectomerus
|
+-- Pleurobema
|
+-- Unionini
Rosenberg et al., 1997 [Publ 2720], fig. 2B iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 17 | 0 | 3 | 18 | 7 | 3 | 33 | +-- Gonideini
|
+-- Pyganodon_grandis
|
+-- Utterbackia
|
+-- Pyganodon_cataracta
|
+-- Lampsilis_straminea
|
+-- Lampsilis_teres
|
+-- Megalonaias **
|
+-- Obliquaria
|
--+-- Theliderma **
|
+-- Quadrula **
|
+-- Uniomerus **
|
+-- Amblema
|
+-- Elliptio
|
+-- Fusconaia
|
+-- Plectomerus
|
+-- Aculamprotulina
|
+-- Unionina
Rosenberg et al., 1994 [Publ 2719], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 10 | 0 | 3 | 15 | 7 | 3 | 27 | +-- Pyganodon_cataracta
|
+-- Pyganodon_grandis
|
+-- Utterbackia
|
+-- Lampsilis_teres
|
+-- Lampsilis_straminea
--+
+-- Megalonaias **
|
+-- Obliquaria
|
+-- Theliderma **
|
+-- Quadrula **
|
+-- Uniomerus **
Huang et al., 2019 [Publ 3843], fig. S3 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 4 | 2 | 3 | 195 | 21 | 3 | 19 | +----- Quadrulina **
+-----------------------------+
| | +-- Megalonaias **
| +--+
--+ +-- Uniomerus **
|
| +-------------------------------- Pleurobemini
| |
+--+ +----------------------------- Amblema
| |
+--+ +-------------------------- Toxolasma
| |
+--+ +-- Obliquaria
| +--------------------+
| | +-- Plectomerus
+--+
| +-------------------- Potamilus_leptodon
| |
+--+ +----------------- Potamilus_alatus
| |
| | +-- Medionidus
+--+ +-----------+
| | +-- Ptychobranchus
| |
+--+ +-- Ellipsaria
| +--------+
| | +-- Truncilla
+--+
| +-------- Leaunio
| |
+--+ +-- Venustaconcha
| +--+
| | +-- Cambarunio
+--+
| +-- Ortmanniana
+--+
+-- Lampsilis
Huang et al., 2019 [Publ 3843], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 4 | 2 | 3 | 197 | 21 | 3 | 19 | +----- Quadrulina **
+--------------------+
| | +-- Megalonaias **
| +--+
--+ +-- Uniomerus **
|
| +----------------------- Pleurobemini
| |
+--+ +-------------------- Amblema
| |
| | +----- Toxolasma
+--+ +-----------+
| | | +-- Obliquaria
| | +--+
| | +-- Plectomerus
+--+
| +-------------- Potamilus_leptodon
| |
| | +----------- Potamilus_alatus
+--+ |
| | +-------- Venustaconcha
| | |
+--+ +-------- Leaunio
| |
| | +-- Ellipsaria
| +-----+
+--+ +-- Truncilla
|
| +-- Medionidus
+-----+
| +-- Ptychobranchus
|
| +----- Cambarunio
+--+
| +-- Ortmanniana
+--+
+-- Lampsilis
Pfeiffer et al., 2018 [Publ 3835], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 4 | 2 | 3 | 0 | 0 | 0 | 19 | +----- Quadrulina **
--+
| +-- Uniomerus **
+--+
+-- Megalonaias **
Krebs et al., 2003 [Publ 2691], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 21 | 20 | 3 | 29 | 17 | 3 | 38 | +-- Megalonaias **
+-----------------------+
| +-- Quadrulina **
|
| +----------- Elliptoideus
| +-----------+
--+ | | +-------- Pleurobema
| | +--+
| | | +----- Uniomerus **
| | +--+
| | | +-- Fusconaia
+--+ +--+
| +-- Eurynia
|
| +-------------------- Amblema
| |
| | +----- Toxolasma
| | +-----------+
+--+ | | +-- Plectomerus
| | +--+
| | +-- Obliquaria
| |
+--+ +----- Ptychobranchus
| +--------+
| | | +-- Potamilus_fragilis
| | +--+
| | +-- Medionidus
+--+
| +-- Villosa
| +--------+
| | +-- Ellipsaria
+--+
| +-------- Lampsilis_teres
+--+
| +----- Lampsilis_siliquoidea
+--+
| +-- Potamilus_purpuratus
+--+
+-- Potamilus_alatus
Lydeard et al., 1996 [Publ 2656], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 18 | 14 | 3 | 23 | 14 | 3 | 35 | +-- Megalonaias **
+-----------------------+
| +-- Quadrulina **
|
| +----- Elliptoideus
--+ |
| +-----------------+ +-- Eurynia
| | | |
| | +--+-- Pleurobema
| | |
+--+ +-- Uniomerus **
|
| +-------------------- Amblema
| |
+--+ +-- Plectomerus
| +--------------+
| | +-- Obliquaria
+--+
| +-------------- Potamilus_fragilis
| |
+--+ +----------- Ptychobranchus
| |
+--+ +-------- Medionidus
| |
+--+ +----- Potamilus_purpuratus
| |
+--+----- Lampsilis
|
| +-- Ellipsaria
+--+
+-- Villosa
Lydeard et al., 1996 [Publ 2656], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 18 | 16 | 3 | 24 | 14 | 3 | 35 | +-- Megalonaias **
+-----------+
| +-- Quadrulina **
|
--+ +----- Amblema
| +-----+
| | | +-- Plectomerus
| | +--+
+--+ +-- Lampsilini
|
| +-------- Elliptoideus
+--+
| +----- Uniomerus **
+--+
| +-- Eurynia
+--+
+-- Pleurobema
Huang et al., 2019 [Publ 3843], fig. S4 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 128 | 73 | 3 | 197 | 21 | 3 | 144 | +----------- Chamberlainia
|
+----------- Megalonaias **
|
+----------- Uniomerus **
|
+----------- Quadrulina **
|
+----------- Sinohyriopsis
|
+----------- Lamprotulini
|
+----------- Gonideini
|
+----------- Pseudodontini
|
+----------- Rectidentini
|
| +-------- Amblema
| |
| +-------- Toxolasma
| |
| +-------- Pleurobemini
| |
| | +-- Obliquaria
| +-----+
| | +-- Plectomerus
+--+
| | +----- Ellipsaria
| | |
| | +----- Potamilus_leptodon
| | |
| | +----- Medionidus
| | |
| | +----- Potamilus_alatus
| | |
--+ | +----- Ptychobranchus
| +--+
| +----- Truncilla
| |
| +----- Venustaconcha
| |
| +----- Cambarunio
| |
| +----- Leaunio
| |
| | +-- Ortmanniana
| +--+
| +-- Lampsilis
|
| +-------- Acuticostini
| |
| +-------- Alasmidonta
| |
| +-------- Anodonta_anatina
| |
| +-------- Anemina
| |
| +-------- Anodonta_cygnea
| |
| +-------- Cristaria
| |
| +-------- Brachyanodon
| |
| +-------- Lasmigona
| |
| +-------- Lepidodesmini
| |
| +-------- Pseudanodonta
| |
| +-------- Pyganodon
| |
+--+-------- Simpsonaias
|
+-------- Strophitus
|
+-------- Utterbackia
|
+-------- Schistodesmus
|
+-------- Unionina
|
+-------- Aculamprotulina
|
+-------- Lanceolariini
|
| +-- Pletholophus
+-----+
| +-- Sinanodonta
|
| +----- Cuneopsis_celtiformis
| |
| +----- Tchangsinaia
| |
+--+----- Cuneopsis_rufescens
|
+----- Nodularia
|
| +-- Cuneopsis_szechenyii
+--+
+-- Cuneopsis_heudei
Lydeard et al., 2000 [Publ 2654], fig. 2d iT| tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 0 | 1 | 11 | 9 | 3 | 21 | +-------- Amblemini
--+
| +----- Lampsilini
+--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Lydeard et al., 2000 [Publ 2654], fig. 2b iT| tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 0 | 1 | 11 | 9 | 3 | 21 | +-------- Lampsilini
--+
| +----- Amblemini
+--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Chase et al., 2018 [Publ 3847], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 1 | 2 | 36 | 15 | 3 | 20 | +----- Pleurobemini
--+
| +-- Megalonaias **
+--+
+-- Quadrulina **
Lydeard et al., 2000 [Publ 2654], fig. 2c iT| tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 1 | 1 | 11 | 9 | 3 | 20 | Lydeard et al., 2000 [Publ 2654], fig. 2a iT| tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 1 | 1 | 11 | 9 | 3 | 20 | +-- Quadrulina **
--+
+-- Pleurobemini
Wu et al., 2021 [Publ 4366], fig. 3-BI iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 71 | 13 | 4 | -- | +-- Uniomerus **
+--------+
| +-- Quadrulina **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Pieri et al., 2018 [Publ 3867], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 18 | 17 | 1 | -- | +-- Quadrulina **
--+
+-- Pleurobemini
Amaro et al., 2017 [Publ 3791], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 14 | 4 | 2 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Graf & Cummings, 2006 [Publ 1968], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 45 | 12 | 3 | -- | +-------- Quadrulina **
--+
| +----- Amblemini
+--+
| +-- Pleurobemini
+--+
+-- Lampsilini
Curole & Kocher, 2002 [Publ 2712], fig. 1a-M iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 2 | 1 | 1 | -- | Hoeh et al., 2009 [Publ 2716], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 44 | 11 | 3 | -- | +-- Obliquaria
+--------+
| +-- Quadrulina **
|
--+ +-------- Amblemini
| |
| +-------- Pleurobemini
+--+
| +----- Truncilla
| |
| +----- Ptychobranchus
+--+
| +-- Ortmanniana
| |
| +-- Cambarunio
+--+
+-- Ligumia
|
+-- Epioblasma
Pfeiffer et al., 2019 [Publ 3937], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 55 | 13 | 4 | -- | +-- Quadrulina **
+-----+
| +-- Uniomerus **
|
--+ +-- Pleurobemini
| +--+
| | +-- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Porto-Hannes et al., 2021 [Publ 4227], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 11 | 0 | 1 | 9 | 9 | 3 | 30 | +-------- Quadrula **
|
--+-------- Pustulosa **
|
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Jeratthitikul et al., 2022 [Publ 4260], fig. S1_1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 74 | 7 | 3 | -- | +-- Amblemini
+--+
| +-- Lampsilini
--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Chapman et al., 2008 [Publ 2744], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 19 | 18 | 4 | -- | +-------------- Quadrulina **
--+
| +----------- Pleurobemini
+--+
| +-------- Amblema
+--+
| +----- Plectomerus
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Freeman et al., 2025 [Publ 4443], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 9 | 9 | 4 | -- | +-- Popenaiadini
+-----+
| +-- Lampsilini
--+
| +----- Amblemini
+--+
| +-- Pleurobemini
+--+
+-- Quadrulina **
Jeratthitikul et al., 2022 [Publ 4260], fig. S2_2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 74 | 7 | 3 | -- | +-- Amblemini
+--+
| +-- Lampsilini
--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Chen et al., 2025 [Publ 4448], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 83 | 13 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Wu et al., 2021 [Publ 4366], fig. 3-ML iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 71 | 13 | 4 | -- | +-- Uniomerus **
+--------+
| +-- Quadrulina **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Teiga-Teixeira et al., 2020 [Publ 4173], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 17 | 12 | 4 | -- | +-- Uniomerus **
+--------+
| +-- Quadrulina **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Wu et al., 2024 [Publ 4350], fig. 8b_S2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 89 | 15 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Lopes-Lima et al., 2017 [Publ 3745], fig. 1-2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 79 | 7 | 3 | -- | +-- Amblemini
+--+
| +-- Lampsilini
--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Mejia-Trujillo & Smith, 2022 [Publ 4274], fig. 2-F iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 13 | 13 | 4 | -- | +-------------- Toxolasma
+--+
| | +----------- Popenaiadini
| +--+
| | +-------- Amblemini
| +--+
| | +----- Pleurobemini
--+ +--+
| | +-- Quadrulina **
| +--+
| +-- Uniomerus **
|
| +----------- Potamilus
| |
+-----+ +-- Lampsilis_cardium
| +-----+
| | +-- Lampsilis_ornata
+--+
| +----- Venustaconcha
+--+
| +-- Lampsilis_siliquoidea
+--+
+-- Lampsilis_powellii
Wu et al., 2022 [Publ 4238], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 71 | 13 | 4 | -- | +-- Uniomerus **
+--------+
| +-- Quadrulina **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Campbell & Lydeard, 2026 [Publ 4810], fig. 3_5 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 138 | 16 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Lampsilini
+--+
| +-- Amblemini
+--+
+-- Popenaiadini
Curole & Kocher, 2005 [Publ 2682], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 9 | 7 | 2 | -- | +----------------- Glebula
|
--+ +-- Amblema
| +-----------+
| | +-- Quadrulina **
+--+
| +----------- Plectomerus
+--+
| +-------- Leaunio
+--+
| +----- Lampsilis_straminea
+--+
| +-- Lampsilis_teres
+--+
+-- Potamilus
Wu et al., 2024 [Publ 4350], fig. 8a_S1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 89 | 15 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Huang et al., 2019 [Publ 3843], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 54 | 8 | 2 | -- | +-- Quadrulina **
+--+
| +-- Uniomerus **
--+
| +-- Pleurobemini
+--+
+-- Lampsilini
Wu et al., 2023 [Publ 4339], fig. 4_S5-aa iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 93 | 15 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Bolotov et al., 2017 [Publ 3761], fig. S1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 137 | 7 | 3 | -- | +-- Amblemini
+--+
| +-- Lampsilini
--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Jadhav & Jamkhedkar, 2009 [Publ 3697], fig. 11 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 23 | 11 | 3 | -- | Wu et al., 2023 [Publ 4339], fig. 4_S5-nt iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 93 | 15 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Inoue et al., 2020 [Publ 3941], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 40 | 39 | 3 | -- | +-------- Quadrulina **
--+
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Bolotov et al., 2017 [Publ 3764], fig. S1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 9 | 0 | 1 | 123 | 7 | 3 | 28 | +-- Amblemini
|
+-- Ortmanniana
|
+-- Lampsilis
|
+-- Cambarunio
|
--+-- Elliptio
|
+-- Eurynia
|
+-- Pleurobema
|
+-- Quadrula_quadrula **
|
+-- Quadrula_verrucosa **
Curole & Kocher, 2002 [Publ 2712], fig. 1b-M iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 2 | 1 | 1 | -- | Chapman et al., 2008 [Publ 2667], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 19 | 18 | 4 | -- | +-------------- Quadrulina **
--+
| +----------- Pleurobemini
+--+
| +-------- Amblema
+--+
| +----- Plectomerus
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Curole & Kocher, 2002 [Publ 2712], fig. 1-F iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 2 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Burlakova et al., 2019 [Publ 3899], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 31 | 31 | 4 | -- | +-- Quadrulina **
+-----------------------+
| +-- Uniomerus **
|
| +----------------------- Plectomerus
--+ |
| | +-------------------- Popenaiadini
| | |
| | +-------------------- Pleurobemini
+--+ |
| | +-- Cyrtonaias
| | +--+
| | | +-- Glebula
| +--------------+
| | | +-- Reginaia
+--+ +--+
| +-- Obliquaria
|
| +----------------- Toxolasma
| |
| | +-------------- Potamilus
| | |
| | | +-- Ellipsaria
| | +-----------+
+--+ | +-- Truncilla
| |
| | +----- Medionidus
| +--------+
| | | +-- Lemiox
| | +--+
+--+ +-- Dromus
|
| +----------- Cambarunio
| |
| +----------- Leaunio
| |
| +----------- Villosa
| |
| | +-- Sagittunio
| +--------+
+--+ +-- Lampsilis_ovata
|
| +----- Venustaconcha
+-----+
| | +-- Epioblasma
| +--+
| +-- Obovaria
|
| +-------- Lampsilis_teres
+--+
| +----- Lampsilis_siliquoidea
+--+
| +-- Lampsilis_hydiana
+--+
+-- Lampsilis_straminea
Hoeh et al., 2009 [Publ 2716], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 45 | 12 | 3 | -- | +-------- Amblemini
+-----------+
| | +----- Pleurobemini
| +--+
| | +-- Obliquaria
--+ +--+
| +-- Quadrulina **
|
| +----------------- Ptychobranchus
+--+
| +-------------- Truncilla
+--+
| +----------- Ortmanniana
+--+
| +-------- Lampsilis
+--+
| +----- Cambarunio
+--+
| +-- Ligumia
+--+
+-- Epioblasma
Dai et al., 2024 [Publ 4358], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 51 | 10 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Huang et al., 2002 [Publ 2657], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 14 | 0 | 2 | 32 | 12 | 3 | 33 | +----------------------------- Megalonaias **
|
| +----- Elliptoideus
--+ +--------------------+
| | | +-- Uniomerus **
| | +--+
+--+ +-- Pleurobema
|
| +----------------------- Amblema
| |
+--+ +-- Obliquaria
| +-----------------+
| | +-- Plectomerus
+--+
| +----------------- Medionidus
+--+
| +-------------- Potamilus_fragilis
+--+
| +----------- Ptychobranchus
+--+
| +-------- Potamilus_purpuratus
+--+
| +----- Lampsilis
+--+
| +-- Villosa
+--+
+-- Ellipsaria
Hoeh et al., 2009 [Publ 2716], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 44 | 11 | 2 | -- | +----- Pleurobemini
+--------------+
| | +-- Obliquaria
| +--+
--+ +-- Quadrulina **
|
| +----------------- Ptychobranchus
+--+
| +-------------- Truncilla
+--+
| +----------- Ortmanniana
+--+
| +-------- Lampsilis
+--+
| +----- Cambarunio
+--+
| +-- Ligumia
+--+
+-- Epioblasma
Pfeiffer et al., 2019 [Publ 3937], fig. 4-5 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 55 | 13 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Huang et al., 2002 [Publ 2657], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 14 | 0 | 2 | 32 | 12 | 3 | 33 | +----------- Megalonaias **
|
| +----- Elliptoideus
| +--+
--+ | | +-- Uniomerus **
| | +--+
| | +-- Pleurobema
| |
| | +----- Potamilus_fragilis
+--+ |
| +----- Medionidus
| |
| +----- Amblema
| |
| | +-- Obliquaria
+--+--+
| +-- Plectomerus
|
| +-- Villosa
| |
| +-- Ellipsaria
| |
+--+-- Lampsilis
|
+-- Potamilus_purpuratus
|
+-- Ptychobranchus
Curole & Kocher, 2005 [Publ 2682], fig. 1-M iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 10 | 7 | 2 | -- | +-------- Quadrulina **
--+
| +----- Plectomerus
+--+
| +-- Amblema
+--+
+-- Lampsilini
Freeman et al., 2025 [Publ 4443], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 10 | 10 | 4 | -- | +-- Popenaiadini
+-----+
| +-- Lampsilini
--+
| +----- Amblemini
+--+
| +-- Pleurobemini
+--+
+-- Quadrulina **
Porto-Hannes et al., 2021 [Publ 4227], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 15 | 0 | 1 | 13 | 13 | 3 | 34 | +-------- Quadrula_quadrula **
|
--+-------- Quadrula_verrucosa **
|
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Freeman et al., 2025 [Publ 4443], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 10 | 10 | 4 | -- | +----- Amblemini
+--+
| | +-- Pleurobemini
| +--+
| +-- Quadrulina **
--+
| +----- Obliquaria
| |
| +----- Toxolasma
+--+
+----- Popenaiadini
|
| +-- Glebula
+--+
+-- Cyrtonaias
Inoue et al., 2020 [Publ 3941], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 39 | 38 | 3 | -- | +----------- Quadrulina **
--+
| +-------- Reginaia
+--+
| +----- Pleurobemini
+--+
| +-- Amblema
+--+
+-- Lampsilini
Dai et al., 2025 [Publ 4440], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 85 | 14 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Smith, 2021 [Publ 4331], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 18 | 7 | 3 | -- | +-- Uniomerus **
+-----+
| +-- Megalonaias **
--+
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Dai et al., 2023 [Publ 4285], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 51 | 13 | 4 | -- | +-- Quadrulina **
+--------+
| +-- Uniomerus **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Wu et al., 2024 [Publ 4388], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 51 | 8 | 3 | -- | +-- Quadrulina **
+-----+
| +-- Uniomerus **
--+
| +----- Pleurobemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Bolotov et al., 2017 [Publ 3761], fig. S2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 1 | 137 | 7 | 3 | -- | +-- Amblemini
+--+
| +-- Lampsilini
--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Fukata & Iigo, 2020 [Publ 4106], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 52 | 9 | 2 | -- | +-- Quadrulina **
+--+
| +-- Uniomerus **
--+
| +-- Pleurobemini
+--+
+-- Lampsilini
Fonseca et al., 2016 [Publ 3775], fig. 1-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 8 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Hou et al., 2025 [Publ 4445], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 51 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Kandl et al., 2001 [Publ 3384], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 8 | 8 | 2 | -- | +-- Amblemini
|
--+-- Quadrulina **
|
+-- Pleurobemini
Lopes-Lima et al., 2020 [Publ 4080], fig. 2-4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 121 | 3 | 3 | -- | +-- Amblemini
|
+-- Pleurobemini
--+
+-- Lampsilini
|
+-- Quadrulina **
Smith et al., 2020 [Publ 4136], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 19 | 19 | 1 | -- | +-- Quadrulina **
--+
+-- Pleurobemini
Guerra et al., 2017 [Publ 3772], fig. 2a iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 24 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Froufe et al., 2019 [Publ 3936], fig. 3-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 18 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Lopes-Lima et al., 2017 [Publ 3792], fig. 2-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 9 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Campbell et al., 2008 [Publ 2675], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 24 | 24 | 3 | -- | +----------- Uniomerus **
|
+----------- Amblemini
|
| +-- Elliptio_arctata
| +-----+
| | +-- Elliptio_crassidens
| |
| | +----- Pleurobema_decisum
--+ | |
| | +----- Pleurobema_rubellum
| | |
| | +----- Pleurobema_georgianum__2
| | |
| | +----- Pleurobema_stabile
| | |
| | +----- Pleurobema_pyriforme
| | |
+--+ +----- Pleurobema_oviforme__2
| |
| +----- Pleurobema_hanleyianum__2
| |
| +----- Pleuronaia
| |
| +----- Pleurobema_georgianum__3
| |
| +----- Pleurobema_cordatum
| |
| +----- Pleurobema_beadleianum
| |
| +----- Hemistena
+--+
+----- Fusconaia_flava
|
+----- Fusconaia_cerina
|
+----- Eurynia
|
+----- Elliptio_arca
|
+----- Lampsilini
|
| +-- Plethobasus
+--+
| +-- Parvaspina
|
| +-- Pleurobema_sintoxia
| |
+--+-- Pleurobema_oviforme__1
| |
| +-- Pleurobema_clava
|
| +-- Pleurobema_athearni
| |
+--+-- Pleurobema_georgianum__1
|
+-- Pleurobema_hanleyianum__1
Hoeh et al., 2002 [Publ 2653], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 10 | 3 | -- | +-- Glebula
+--------+
| +-- Cyrtonaias
|
--+ +-------- Toxolasma
| |
| | +----- Potamilus
| +--+
+--+ | +-- Ligumia
| +--+
| +-- Ortmanniana
|
| +----- Amblemini
+--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Huang et al., 2013 [Publ 3629], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 15 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wang et al., 2018 [Publ 3846], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Dai et al., 2025 [Publ 4440], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 43 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Wei et al., 2016 [Publ 3665], fig. 0 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 5 | 1 | 1 | -- | Mulvey et al., 1997 [Publ 3390], fig. 1_3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 4 | 3 | 1 | -- | +-- Megalonaias **
--+
+-- Amblemini
Reis et al., 2013 [Publ 3580], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 13 | 1 | 1 | -- | +-- Amblemini
--+
+-- Quadrulina **
Xue et al., 2016 [Publ 3659], fig. 1-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 1 | 1 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Froufe et al., 2015 [Publ 3664], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 11 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Chen et al., 2023 [Publ 4302], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Guerra et al., 2019 [Publ 4168], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Froufe et al., 2019 [Publ 3936], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 43 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wu et al., 2018 [Publ 3836], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 52 | 8 | 3 | -- | Wen et al., 2017 [Publ 3795], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 15 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Chase et al., 2018 [Publ 3861], fig. 1-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 10 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Amaro et al., 2017 [Publ 3791], fig. 3A-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 6 | 1 | 1 | -- | Reis et al., 2013 [Publ 3580], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 14 | 1 | 1 | -- | +-- Amblemini
--+
+-- Quadrulina **
Chen et al., 2023 [Publ 4302], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Soroka & Burzynski, 2018 [Publ 3762], fig. 3-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 6 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Roe & Hoeh, 2003 [Publ 2655], fig. 4f3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 10 | 3 | -- | +-------- Amblemini
--+
| +----- Lampsilini
+--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Huang et al., 2019 [Publ 3843], fig. 3-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 22 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wu et al., 2023 [Publ 4339], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 18 | 5 | 2 | -- | +----- Lampsilini
--+
| +-- Pleurobemini
+--+
+-- Quadrulina **
Whelan et al., 2011 [Publ 3380], fig. 1-ML iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 35 | 4 | 3 | -- | +-------- Pleurobemini
--+
| +----- Quadrulina **
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Chen et al., 2025 [Publ 4487], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 1 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Plazzi et al., 2011 [Publ 3676], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 17 | 4 | 2 | -- | +-- Quadrulina **
|
--+-- Pleurobemini
|
+-- Lampsilini
Wu et al., 2019 [Publ 3864], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 45 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Zieritz et al., 2021 [Publ 4231], fig. 3-F_5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 47 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wu et al., 2016 [Publ 3711], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 20 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Pavan-Kumar et al., 2022 [Publ 4263], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 36 | 6 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Guerra et al., 2017 [Publ 3772], fig. 2b-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 7 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Inoue et al., 2014 [Publ 3635], fig. 3A iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 19 | 3 | 3 | -- | Krebs, 2004 [Publ 2692], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 4 | 2 | 2 | -- | Amaro et al., 2017 [Publ 3791], fig. 2-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 9 | 3 | 2 | -- | +----- Lampsilini
--+
| +-- Pleurobemini
+--+
+-- Quadrulina **
Burzynski et al., 2017 [Publ 3774], fig. 1-bottom iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 12 | 1 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wu et al., 2020 [Publ 4121], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 47 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Pavan-Kumar et al., 2022 [Publ 4263], fig. S5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 38 | 6 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Sheng et al., 2014 [Publ 3704], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 7 | 2 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Graf & Ó Foighil, 2000 [Publ 2648], fig. 2B iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 29 | 16 | 3 | -- | +-- Quadrulina **
+--+
| +-- Pleurobemini
--+
| +-- Amblemini
+--+
+-- Lampsilini
Campbell & Lydeard, 2026 [Publ 4810], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 46 | 6 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Chen et al., 2025 [Publ 4487], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Hauck et al., 2023 [Publ 4333], fig. 9 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 12 | 12 | 3 | -- | +-------- Quadrulina **
--+
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Froufe et al., 2019 [Publ 3936], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 18 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Mejia-Trujillo & Smith, 2022 [Publ 4274], fig. 2-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 5 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Guerra et al., 2019 [Publ 4168], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Perkins et al., 2017 [Publ 3777], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 23 | 23 | 2 | -- | +-- Amblemini
|
--+-- Uniomerus **
|
+-- Pleurobemini
Amaro et al., 2017 [Publ 3791], fig. 3B iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 11 | 2 | 1 | -- | Inoue et al., 2020 [Publ 4128], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 22 | 22 | 3 | -- | +----------- Quadrulina **
--+
| +-------- Reginaia
+--+
| +----- Amblema
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Klymus et al., 2025 [Publ 4814], fig. 3B iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 35 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Feng et al., 2016 [Publ 3800], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 13 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Graf & Ó Foighil, 2000 [Publ 2648], fig. 2A iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 29 | 16 | 3 | -- | +-- Quadrulina **
+--+
| +-- Pleurobemini
--+
| +-- Amblemini
+--+
+-- Lampsilini
Hou et al., 2024 [Publ 4400], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 74 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Froufe et al., 2019 [Publ 3936], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 43 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Bolotov et al., 2018 [Publ 3826], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 81 | 6 | 3 | -- | Amaro et al., 2017 [Publ 3791], fig. 3B-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 6 | 1 | 1 | -- | Guerra et al., 2019 [Publ 4168], fig. 2-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 21 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Fonseca et al., 2016 [Publ 3775], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Froufe et al., 2015 [Publ 3664], fig. 1-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 7 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Bogan & Hoeh, 2000 [Publ 2650], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 33 | 10 | 3 | -- | +-------- Toxolasma
|
+-------- Amblemini
|
| +-- Glebula
+-----+
--+ +-- Cyrtonaias
|
| +-- Quadrulina **
+-----+
| +-- Pleurobemini
|
| +----- Potamilus
+--+
| +-- Ortmanniana
+--+
+-- Ligumia
Guerra et al., 2019 [Publ 4168], fig. 1-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 21 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wu et al., 2021 [Publ 4366], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 67 | 11 | 4 | -- | +----------- Quadrulina **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Hoeh et al., 2001 [Publ 2652], fig. 14f4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 10 | 3 | -- | +-------- Amblemini
--+
| +----- Lampsilini
+--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Inoue et al., 2020 [Publ 4128], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 21 | 21 | 3 | -- | +----------- Quadrulina **
--+
| +-------- Reginaia
+--+
| +----- Amblema
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Smith et al., 2019 [Publ 3951], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 40 | 3 | -- | +-------------------------------------- Quadrulina **
|
| +-- Amblema
--+ +--------------------------------+
| | +-- Reginaia
| |
+--+ +-------------------------------- Plectomerus
| |
| | +----------------------------- Toxolasma
+--+ |
| | +----- Popenaiadini
| | +--------------------+
+--+ | | +-- Glebula
| | +--+
| | +-- Cyrtonaias
| |
+--+ +----------------------- Obliquaria
| |
| | +-- Medionidus
| | +--+
| | | +-- Lemiox
+--+ +--------------+
| | | +-- Cyprogenia
| | +--+
| | +-- Ptychobranchus
| |
+--+ +-------- Potamilus
| +--------+
| | | +----- Atlanticoncha
| | +--+
| | | +-- Ellipsaria
| | +--+
+--+ +-- Truncilla
|
| +-- Hamiota
| +-----------+
| | +-- Villosa
| |
+--+ +----------- Ligumia
| |
| | +----- Venustaconcha
+--+ +--+
| | | +-- Obovaria
| | +--+
+--+ +-- Epioblasma
|
| +----- Ortmanniana_pectorosa
+--+
| +-- Ortmanniana_ligamentina
+--+
+-- Lampsilis
Huang et al., 2018 [Publ 3828], fig. 6-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 16 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wu et al., 2021 [Publ 4366], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 90 | 5 | 2 | -- | +----- Lampsilini
--+
| +-- Quadrulina **
+--+
+-- Pleurobemini
Wu et al., 2021 [Publ 4366], fig. 2_I iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 90 | 5 | 2 | -- | +----- Quadrulina **
--+
| +-- Pleurobemini
+--+
+-- Lampsilini
Lopes-Lima et al., 2017 [Publ 3792], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 31 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Campbell et al., 2008 [Publ 2675], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 24 | 24 | 3 | -- | +----------------------- Uniomerus **
|
+----------------------- Amblemini
|
| +-------------------- Lampsilini
| |
| +-------------------- Eurynia
| |
| +-------------------- Fusconaia_cerina
--+ |
| +-------------------- Fusconaia_flava
| |
| +-------------------- Hemistena
| |
| +-------------------- Plethobasus
| |
| +-------------------- Parvaspina
| |
+--+-------------------- Pleurobema_cordatum
|
+-------------------- Pleuronaia
|
+-------------------- Pleurobema_oviforme__2
|
+-------------------- Pleurobema_stabile
|
+-------------------- Elliptio
|
| +----------------- Pleurobema_decisum
| |
| +----------------- Pleurobema_sintoxia
| |
| +----------------- Pleurobema_clava
+--+
+----------------- Pleurobema_oviforme__1
|
| +-------------- Pleurobema_beadleianum
| |
+--+ +----------- Pleurobema_pyriforme
| |
+--+ +-------- Pleurobema_rubellum
| |
| +-------- Pleurobema_hanleyianum__2
+--+
+-------- Pleurobema_georgianum__2
|
| +----- Pleurobema_hanleyianum__1
+--+
| +-- Pleurobema_georgianum__1
+--+
+-- Pleurobema_athearni
Guerra et al., 2017 [Publ 3772], fig. 2b iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 24 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Soroka & Burzynski, 2018 [Publ 3762], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 6 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Roe & Hoeh, 2003 [Publ 2655], fig. 4f2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 10 | 3 | -- | Turner et al., 2000 [Publ 2722], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 13 | 11 | 3 | -- | Hoeh et al., 2001 [Publ 2652], fig. 14f2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 10 | 3 | -- | Wu et al., 2020 [Publ 4147], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 29 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Burzynski et al., 2017 [Publ 3774], fig. 1-top iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 12 | 1 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Dai et al., 2025 [Publ 4440], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 43 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Inoue et al., 2013 [Publ 3528], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 16 | 14 | 3 | -- | +-- Pleurobemini
+-----------+
| +-- Quadrulina **
|
--+ +-- Paetulunio
| +--------+
| | +-- Amblemini
+--+
| +-------- Lampsilis
| |
+--+ +----- Leaunio
| |
+--+----- Obovaria
|
| +-- Villosa
+--+
+-- Cambarunio
Chase et al., 2018 [Publ 3861], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 10 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wen et al., 2017 [Publ 3795], fig. 3-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 5 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Cha et al., 2018 [Publ 3916], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 37 | 4 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Huang et al., 2018 [Publ 3828], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 41 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Aunins et al., 2018 [Publ 3863], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 39 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wu et al., 2018 [Publ 4143], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 21 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Pfeiffer & Graf, 2013 [Publ 3531], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 4 | 3 | -- | +-------- Quadrulina **
--+
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Lampsilini
Smith et al., 2020 [Publ 4129], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 27 | 27 | 4 | -- | +----------- Quadrulina **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Whelan et al., 2011 [Publ 3380], fig. 1-MP iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 35 | 4 | 3 | -- | +----- Quadrulina **
|
--+----- Pleurobemini
|
| +-- Amblemini
+--+
+-- Lampsilini
Smith et al., 2020 [Publ 4129], fig. 2-3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 27 | 27 | 4 | -- | +----------- Quadrulina **
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaiadini
+--+
+-- Lampsilini
Zhou et al., 2017 [Publ 3788], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 15 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Wang et al., 2018 [Publ 3865], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Huang et al., 2013 [Publ 3629], fig. 3-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 5 | 1 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Dai et al., 2024 [Publ 4386], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Zhou et al., 2016 [Publ 3710], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 19 | 3 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Guerra et al., 2017 [Publ 3772], fig. 2a-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 7 | 1 | 1 | -- | +-- Lampsilini
--+
+-- Quadrulina **
Chen et al., 2012 [Publ 3624], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 6 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Kandl et al., 2001 [Publ 3384], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 5 | 5 | 1 | -- | +-- Elliptio
|
--+-- Uniomerus **
|
+-- Pleurobema
Burzynski & Soroka, 2018 [Publ 3862], fig. 5a iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 19 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Amaro et al., 2017 [Publ 3791], fig. 3A iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 11 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Chen et al., 2020 [Publ 4170], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 33 | 6 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Xiang et al., 2025 [Publ 4433], fig. 7 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 12 | 0 | 0 | -- | Klymus et al., 2025 [Publ 4814], fig. 3A iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 35 | 5 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
Pfeiffer & Graf, 2015 [Publ 3620], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 39 | 4 | 2 | -- | +----- Quadrulina **
--+
| +-- Amblemini
+--+
+-- Lampsilini
Bolotov et al., 2020 [Publ 4084], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 35 | 6 | 3 | -- | +-- Amblemini
|
+-- Ortmanniana
|
+-- Lampsilis
|
--+-- Cambarunio
|
+-- Elliptio
|
+-- Pleurobema
|
+-- Quadrulina **
Zieritz et al., 2021 [Publ 4231], fig. 3-M_4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 22 | 2 | 1 | -- | +-- Quadrulina **
--+
+-- Lampsilini
|
