The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)
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Bivalvia | Unionoida | Unionoidea | Unionidae | Ambleminae tribe POPENAIADINI Heard & Guckert, 1970: cladomics
6 genera Cladomics Summarynumber of species = 36 (36 total)
number of subtaxa (rank: genus) = 6
analyzed in 43 cladograms in 27 publications
maximum number of terminals in an analysis =
5 (mean = 1.47, median = 1)
maximum number of subtaxa (genus) =
3 (mean = 1.28, median = 1) Basic Monophylynumber of cladograms that test basic monophyly
(i.e., more than 1 included species) = 11
maximum number of included terminals = 5, (mean =
2.82 median = 2)
maximum number of subtaxa (genus) =
3 (mean = 2.09, median = 2)
proportion of cladograms that support basic monophyly
= 1 Rigorous Monophylynumber of cladograms that test rigorous monophyly = 11
(Rigorous monophyly refers to cladograms with terminal species representing
more than one subtaxon (genus) taxon and at least one rigorous outgroup (family).
maximum number of included terminals = 5 (mean = 2.82, median = 2)
maximum number of subtaxa (genus) =
3 (mean = 2.09, median = 2)
proportion of cladograms that support rigorous monophyly = 1
Pairwise Cladogram ClusteringThe graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.
Freeman et al., 2025 [Publ 4443], fig. 2 ** Top Monophyly Cladogram ** iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 4 | 3 | 3 | 2 | 2 | 1 | 0 | +-------- Lampsilini
--+
| +----- Popenaias **
+--+
| +-- Nephronaias **
+--+
+-- Psoronaias **
Campbell et al., 2005 [Publ 2660], fig. 1 ** Top Sister-Group Cladogram ** iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 101 | 89 | 4 | -- | +-------------- Quadrulini
|
--+ +----------- Pleurobemini
| |
+--+ +-- Plectomerus
| +-----+
| | +-- Reginaia
+--+
| +----- Lampsilini
+--+
| +-- Popenaias **
+--+
+-- Amblema
Inoue et al., 2020 [Publ 4128], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 5 | 5 | 7 | 2 | 17 | 17 | 3 | 7 | +-------------- Quadrulini
--+
| +----------- Reginaia
+--+
| +-------- Amblema
+--+
| +----- Lampsilini
+--+
| +-- Popenaias **
+--+
+-- Psoronaias **
Inoue et al., 2020 [Publ 4128], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 5 | 5 | 3 | 2 | 18 | 18 | 3 | 8 | +-------------- Quadrulini
--+
| +----------- Reginaia
+--+
| +-------- Amblema
+--+
| +----- Lampsilini
+--+
| +-- Popenaias **
+--+
+-- Psoronaias **
Pfeiffer et al., 2019 [Publ 3866], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 2 | 2 | 28 | 27 | 4 | 3 | +-------------- Quadrulini
--+
| +----------- Pleurobemini
+--+
| +-------- Amblemini
+--+
| +----- Lampsilini
+--+
| +-- Popenaias **
+--+
+-- Psoronaias **
Smith et al., 2020 [Publ 4129], fig. 2-3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 26 | 26 | 4 | -- | +-------------- Quadrulini
--+
| +----------- Pleurobemini
+--+
| +-------- Amblemini
+--+
| +----- Lampsilini
+--+
| +-- Popenaias **
+--+
+-- Psoronaias **
Freeman et al., 2025 [Publ 4443], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 9 | 9 | 4 | -- | +----- Lampsilini
+--+
| | +-- Nephronaias **
| +--+
--+ +-- Popenaias **
|
| +----- Amblemini
+--+
| +-- Pleurobemini
+--+
+-- Quadrulini
Smith et al., 2020 [Publ 4129], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 26 | 26 | 4 | -- | +-------------- Quadrulini
--+
| +----------- Pleurobemini
+--+
| +-------- Amblemini
+--+
| +----- Lampsilini
+--+
| +-- Popenaias **
+--+
+-- Psoronaias **
Freeman et al., 2025 [Publ 4443], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 10 | 10 | 4 | -- | +----- Lampsilini
+--+
| | +-- Nephronaias **
| +--+
--+ +-- Popenaias **
|
| +----- Amblemini
+--+
| +-- Pleurobemini
+--+
+-- Quadrulini
Pfeiffer et al., 2019 [Publ 3937], fig. 4-5 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 55 | 13 | 4 | -- | +-------------- Quadrulini
--+
| +----------- Pleurobemini
+--+
| +-------- Amblemini
+--+
| +----- Lampsilini
+--+
| +-- Psoronaias **
+--+
+-- Popenaias **
Freeman et al., 2025 [Publ 4443], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 10 | 10 | 4 | -- | +----- Amblemini
+--+
| | +-- Pleurobemini
| +--+
| +-- Quadrulini
--+
| +----- Obliquaria
| |
| +----- Toxolasma
| |
+--+ +-- Glebula
+--+
| +-- Cyrtonaias
|
| +-- Nephronaias **
+--+
+-- Popenaias **
Pfeiffer et al., 2019 [Publ 3937], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 55 | 13 | 4 | -- | +----------- Quadrulini
|
--+ +-- Pleurobemini
| +-----+
| | +-- Amblemini
+--+
| +----- Lampsilini
+--+
| +-- Psoronaias **
+--+
+-- Popenaias **
Chen et al., 2025 [Publ 4448], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 85 | 14 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Wu et al., 2024 [Publ 4350], fig. 8b_S2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 91 | 16 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Campbell & Lydeard, 2012 [Publ 3308], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 41 | 41 | 4 | -- | +-------------------- Quadrulini
|
--+ +----------------- Pleurobemini
| |
+--+ +-- Reginaia
| +-----------+
| | +-- Amblema
+--+
| +----------- Popenaias **
| |
+--+ +-------- Cyrtonaias
| |
+--+-------- Plectomerus
|
| +----- Lemiox
+--+
| +-- Lampsilis
+--+
+-- Obovaria
Campbell & Lydeard, 2012 [Publ 3308], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 42 | 42 | 4 | -- | +-------------------------------- Quadrulini
|
| +-------------- Elliptoideus
| +--------------+
| | | +----------- Plectomerus
--+ | +--+
| | | +-------- Obovaria_unicolor
| | +--+
| | | +----- Obovaria_olivaria
| | +--+
| | | +-- Lampsilis
+--+ +--+
| +-- Lemiox
|
| +-------- Cyrtonaias
| +-----------------+
| | | +----- Reginaia
| | +--+
| | | +-- Popenaias **
+--+ +--+
| +-- Amblema
|
| +----------------------- Elliptio
| |
+--+ +-------------------- Eurynia
| |
| | +----------------- Parvaspina
+--+ |
| | +-- Hemistena
| +--------------+
+--+ +-- Plethobasus
|
| +-------------- Pleuronaia
| |
| | +-- Pleurobema_cordatum
+--+ +--------+
| | +-- Fusconaia
| |
+--+ +-- Pleurobema_stabile
| +-----+
| | +-- Pleurobema_sintoxia
+--+
| +----- Pleurobema_decisum
+--+
| +-- Pleurobema_clava
+--+
+-- Pleurobema_pyriforme
Wu et al., 2024 [Publ 4388], fig. 6 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 53 | 9 | 3 | -- | +-------- Quadrulini
--+
| +----- Pleurobemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Wu et al., 2023 [Publ 4339], fig. 4_S5-nt iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 95 | 16 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Chapman et al., 2008 [Publ 2744], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 20 | 19 | 4 | -- | +-------------- Quadrulini
--+
| +----------- Pleurobemini
+--+
| +-------- Amblema
+--+
| +----- Plectomerus
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Mejia-Trujillo & Smith, 2022 [Publ 4274], fig. 2-F iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 14 | 14 | 4 | -- | +----------- Toxolasma
+--+
| | +-------- Popenaias **
| +--+
| | +----- Amblemini
| +--+
--+ | +-- Quadrulini
| +--+
| +-- Pleurobemini
|
| +----------- Potamilus
| |
+--+ +-- Lampsilis_cardium
| +-----+
| | +-- Lampsilis_ornata
+--+
| +----- Venustaconcha
+--+
| +-- Lampsilis_siliquoidea
+--+
+-- Lampsilis_powellii
Teiga-Teixeira et al., 2020 [Publ 4173], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 18 | 13 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Zanatta & Murphy, 2006 [Publ 2672], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 55 | 53 | 3 | -- | +-------- Lampsilini
--+
| +----- Pleurobemini
+--+
| +-- Amblemini
+--+
+-- Popenaias **
Wu et al., 2021 [Publ 4366], fig. 3-ML iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 73 | 14 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Dai et al., 2024 [Publ 4358], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 52 | 11 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Campbell et al., 2005 [Publ 2660], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 101 | 89 | 4 | -- | +-------------------------- Quadrulini
|
| +----------------------- Pleurobemini
--+ |
| | +----- Popenaias **
| | +--------------+
| | | | +-- 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
Campbell & Lydeard, 2012 [Publ 3308], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 41 | 41 | 4 | -- | +-------------------- Quadrulini
|
--+ +----------------- Pleurobemini
| |
| | +-- Popenaias **
+--+ +-----------+
| | +-- Amblema
| |
+--+ +-- Cyrtonaias
| +--------+
| | +-- Reginaia
+--+
| +-------- Plectomerus
+--+
| +----- Lemiox
+--+
| +-- Lampsilis
+--+
+-- Obovaria
Wu et al., 2022 [Publ 4238], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 73 | 14 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Campbell & Lydeard, 2012 [Publ 3309], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 72 | 72 | 4 | -- | +----------------------------------- Quadrulini
|
--+ +-------------------------------- Pleurobemini
| |
| | +-- Amblema
+--+ +--------------------------+
| | +-- Reginaia
| |
+--+ +-------------------------- Popenaias **
| |
| | +-- Glebula
+--+ +--------------------+
| | +-- Paetulunio
| |
+--+ +-- Toxolasma
| +-----------------+
| | +-- Plectomerus
| |
+--+ +----------------- Obliquaria
| |
| | +----- Potamilus
+--+ +--------+
| | | +-- Dromus
| | +--+
+--+ +-- Lemiox
|
| +----------- Lampsilis_ovata
| |
+--+ +-------- Lampsilis_virescens
| |
+--+ +-- Epioblasma
| +--+
| | +-- Obovaria_arkansasensis
+--+
| +-- Obovaria_olivaria
+--+
+-- Venustaconcha
Burlakova et al., 2012 [Publ 3486], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 40 | 3 | -- | +----------- Amblema
|
--+ +-- Lampsilis
| +-----+
| | +-- Plectomerus
+--+
| +----- Popenaias **
+--+
| +-- Cyrtonaias
+--+
+-- Pleurobemini
Chapman et al., 2008 [Publ 2667], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 20 | 19 | 4 | -- | +-------------- Quadrulini
--+
| +----------- Pleurobemini
+--+
| +-------- Amblema
+--+
| +----- Plectomerus
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Dai et al., 2025 [Publ 4440], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 87 | 15 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Campbell & Lydeard, 2012 [Publ 3308], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 41 | 41 | 4 | -- | +----------------- Quadrulini
|
--+ +-------------- Pleurobemini
| |
| | +-- Popenaias **
+--+ +--------+
| | +-- Amblema
| |
+--+ +-------- Plectomerus
| |
| | +-- Cyrtonaias
+--+-----+
| +-- Reginaia
|
| +----- Lemiox
+--+
| +-- Lampsilis
+--+
+-- Obovaria
Inoue et al., 2018 [Publ 3858], fig. 1_3a-c iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 103 | 81 | 4 | -- | +-------- Reginaia
|
+-------- Amblema
|
| +-- Quadrulini
+-----+
| +-- Pleurobemini
--+
| +----- Popenaias **
| |
| | +-- Dromus
| | |
| | +-- Epioblasma
| | |
+--+ +-- Glebula
| |
| +-- Lampsilis_ovata
| |
| +-- Lampsilis_virescens
| |
| +-- Lemiox
+--+
+-- Obliquaria
|
+-- Obovaria_olivaria
|
+-- Obovaria_unicolor
|
+-- Plectomerus
|
+-- Potamilus_alatus
|
+-- Potamilus_capax
Wu et al., 2021 [Publ 4366], fig. 3-BI iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 73 | 14 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Wu et al., 2024 [Publ 4350], fig. 8a_S1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 91 | 16 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Wu et al., 2021 [Publ 4366], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 68 | 11 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Burlakova et al., 2019 [Publ 3899], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 32 | 32 | 4 | -- | +-------------------------- Quadrulini
|
| +----------------------- Plectomerus
--+ |
| | +-------------------- Popenaias **
| | |
| | +-------------------- 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
Wu et al., 2023 [Publ 4339], fig. 4_S5-aa iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 95 | 16 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Dai et al., 2023 [Publ 4285], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 52 | 14 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Popenaias **
+--+
+-- Lampsilini
Smith et al., 2019 [Publ 3951], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 40 | 3 | -- | +-------------------------------------- Quadrulini
|
| +-- Amblema
--+ +--------------------------------+
| | +-- Reginaia
| |
+--+ +-------------------------------- Plectomerus
| |
| | +----------------------------- Toxolasma
+--+ |
| | +----- Popenaias **
| | +--------------------+
+--+ | | +-- Glebula
| | +--+
| | +-- Cyrtonaias
| |
+--+ +----------------------- Obliquaria
| |
| | +-- Medionidus
| | +--+
| | | +-- Lemiox
+--+ +--------------+
| | | +-- Cyprogenia
| | +--+
| | +-- Ptychobranchus
| |
+--+ +-------- Potamilus
| +--------+
| | | +----- Atlanticoncha
| | +--+
| | | +-- Ellipsaria
| | +--+
+--+ +-- Truncilla
|
| +-- Hamiota
| +-----------+
| | +-- Villosa
| |
+--+ +----------- Ligumia
| |
| | +----- Venustaconcha
+--+ +--+
| | | +-- Obovaria
| | +--+
+--+ +-- Epioblasma
|
| +----- Ortmanniana_pectorosa
+--+
| +-- Ortmanniana_ligamentina
+--+
+-- Lampsilis
Neemuchwala et al., 2023 [Publ 4292], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 26 | 25 | 4 | -- | +----------- Quadrulini
--+
| +-------- Pleurobemini
+--+
| +----- Amblemini
+--+
| +-- Lampsilini
+--+
+-- Popenaias **
Zanatta & Murphy, 2006 [Publ 2672], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 56 | 54 | 3 | -- | +----- Pleurobemini
|
--+ +-- Amblemini
| |
+--+-- Popenaias **
|
+-- Lampsilini
Campbell & Lydeard, 2012 [Publ 3309], fig. 1-2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 72 | 72 | 4 | -- | +-------------- Quadrulini
|
--+ +----------- Pleurobemini
| |
+--+ +-- Plectomerus
| +-----+
| | +-- Reginaia
+--+
| +----- Lampsilini
+--+
| +-- Popenaias **
+--+
+-- Amblema
|
