MUSSELpdb
Taxonomy
Species & Genus Richness of
freshwater mussels
by Taxonomy
Species & Genus Richness of
freshwater bivalves
by Family
Geography
Species Richness & Endemism of
freshwater mussels
by Geography
Data last updated
Thu 19 March 2026
Bivalvia | Unionida | Unionoidea | Unionidae | Ambleminae | Lampsilini
genus Villosa Frierson, 1927: cladomics
4 Recent valid species (4 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 = 4 (4 total)
analyzed in 16 cladograms in 12 publications
maximum number of terminals in an analysis = 3 (mean = 1.13, median = 1)
Basic Monophyly
number of cladograms that test basic monophyly (i.e., more than 1 included species) = 1
maximum number of included terminals = 3, (mean = 3 median = 3)
proportion of cladograms that support basic monophyly = 0
Cladograms
Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14
** Top Monophyly Cladogram ** ** Top Sister-Group Cladogram **
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 70 | 78 | 3 | 628 | 90 | 26 | 0 |
+-------- 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
Turner et al., 2000 [Publ 2722], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 13 | 8 | 2 | -- |
+-------------------- Medionidus | --+ +----------------- Lampsilis_teres | | | +----------------- Villosa_delumbis ** +--+ | +-- Lampsilis_satura | +-----------+ | | +-- Lampsilis_cardium +--+ | +----------- Lampsilis_reeveiana | | +--+ +-------- Lampsilis_hydiana__1 | | +--+-------- Lampsilis_hydiana__2 | | +----- Lampsilis_siliquoidea__2 +--+ | +-- Lampsilis_siliquoidea__1 +--+ +-- Lampsilis_powellii
Campbell et al., 2005 [Publ 2660], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 101 | 42 | 22 | -- |
+----- Obliquaria
+--------+
| | +-- Cyrtonaias
| +--+
| +-- Glebula
|
| +----------- Sagittunio
| |
| +----------- Leaunio
| |
| +----------- Potamilus
| |
| +----------- Hamiota
| |
| | +-- Ellipsaria
--+ +--------+
| | +-- Truncilla
| |
| | +----- Lampsilis_teres
| +-----+
| | | +-- Ortmanniana_ligamentina
| | +--+
| | +-- Lampsilis_siliquoidea
| |
| | +----- Villosa_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
Buhay et al., 2002 [Publ 2661], fig. 2A
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 6 | 6 | 3 | -- |
+----- Lampsilis | --+----- Epioblasma | | +-- Cambarunio +--+ +-- Villosa_villosa **
Huang et al., 2002 [Publ 2657], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 33 | 7 | 6 | -- |
+-- Villosa_delumbis ** | +-- Ellipsaria | --+-- Lampsilis | +-- Potamilus | +-- Ptychobranchus
Patterson et al., 2021 [Publ 4222], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 3 | 0 | 0 | -- |
-- Villosa_villosa **
Burlakova et al., 2019 [Publ 3899], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 32 | 25 | 17 | -- |
+----------------- Toxolasma | | +-------------- Potamilus | | | | +-- Ellipsaria | +-----------+ --+ | +-- Truncilla | | | | +----- Medionidus | +--------+ | | | +-- Lemiox | | +--+ +--+ +-- Dromus | | +----------- Cambarunio | | | +----------- Leaunio | | | +----------- Villosa_villosa ** | | | | +-- Sagittunio | +--------+ +--+ +-- Lampsilis_ovata | | +----- Venustaconcha +-----+ | | +-- Epioblasma | +--+ | +-- Obovaria | | +-------- Lampsilis_teres +--+ | +----- Lampsilis_siliquoidea +--+ | +-- Lampsilis_hydiana +--+ +-- Lampsilis_straminea
Buhay et al., 2002 [Publ 2661], fig. 2B
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 6 | 6 | 3 | -- |
+----- Lampsilis | --+----- Epioblasma | | +-- Cambarunio +--+ +-- Villosa_villosa **
Lydeard et al., 1996 [Publ 2656], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 27 | 8 | 6 | -- |
+-------------------- Obliquaria --+ | +----------------- Medionidus +--+ | +-------------- Ptychobranchus +--+ | +----------- Potamilus_fragilis +--+ | +-------- Potamilus_purpuratus +--+ | +----- Ellipsaria +--+ | +-- Villosa_delumbis ** +--+ +-- Lampsilis
Lydeard et al., 1996 [Publ 2656], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 26 | 8 | 6 | -- |
+-------------- Potamilus_fragilis | --+ +----------- Ptychobranchus | | +--+ +-------- Medionidus | | +--+ +----- Potamilus_purpuratus | | +--+----- Lampsilis | | +-- Ellipsaria +--+ +-- Villosa_delumbis **
Smith et al., 2019 [Publ 3951], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 40 | 35 | 19 | -- |
+----------------------- Obliquaria | | +-- Medionidus | +--+ | | +-- Lemiox --+ +--------------+ | | | +-- Cyprogenia | | +--+ | | +-- Ptychobranchus | | +--+ +-------- Potamilus | +--------+ | | | +----- Atlanticoncha | | +--+ | | | +-- Ellipsaria | | +--+ +--+ +-- Truncilla | | +-- Hamiota | +-----------+ | | +-- Villosa_villosa ** | | +--+ +----------- Ligumia | | | | +----- Venustaconcha +--+ +--+ | | | +-- Obovaria | | +--+ +--+ +-- Epioblasma | | +----- Ortmanniana_pectorosa +--+ | +-- Ortmanniana_ligamentina +--+ +-- Lampsilis
Campbell et al., 2005 [Publ 2660], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 101 | 42 | 22 | -- |
+-------- Toxolasma
+--------------------+
| | +----- Obliquaria
| +--+
| | +-- Cyrtonaias
| +--+
| +-- Glebula
|
| +-------------- Potamilus
| |
| | +-------- Medionidus_acutissimus
--+ | +--+
| +-----------+ | | +----- Lemiox
| | | | +--+
| | | | | +-- Dromus
| | | | +--+
| | +--+ +-- Cyprogenia
| | |
| | | +-- Sagittunio
| | | +--+
| | | | +-- Medionidus_conradicus
+--+ +-----+
| | +-- Ptychobranchus
| +--+
| +-- Venustaconcha_ellipsiformis__2
|
| +-- Ellipsaria
| +--------------------+
| | +-- Truncilla
| |
| | +-------------------- Villosa_villosa **
+--+ |
| | +-- Obovaria_olivaria
| | +--------+
| | | +-- Venustaconcha_ellipsiformis__1
| | +-----+
+--+ | | +-------- Epioblasma
| | +--+
| | | +----- Obovaria_unicolor
| | +--+
| | | +-- Obovaria_arkansasensis
| | +--+
| | +-- Obovaria_subrotunda
+--+
| +-------- Hamiota
| |
| +-----+ +-- Lampsilis_cardium
| | | +--+
| | | | +-- Lampsilis_ovata
| | +--+
| | | +-- Ortmanniana_pectorosa
+--+ +--+
| +-- Lampsilis_ornata
|
| +-- Ligumia
| +--------+
| | +-- Cambarunio
+--+
| +-------- Lampsilis_teres
+--+
| +----- Ortmanniana_ligamentina
+--+
| +-- Lampsilis_siliquoidea
+--+
+-- Leaunio
Inoue et al., 2013 [Publ 3528], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 16 | 9 | 5 | -- |
+-------- Lampsilis | --+ +----- Leaunio | | +--+----- Obovaria | | +-- Villosa_villosa ** +--+ +-- Cambarunio
Roe et al., 2002 [Publ 2689], fig. 3a
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 9 | 9 | 4 | -- |
+----- Ligumia | +----- Obliquaria --+ +----- Lampsilis | | +-- Villosa_villosa ** +--+ +-- Hamiota
Huang et al., 2002 [Publ 2657], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 33 | 7 | 6 | -- |
+----------------- Medionidus --+ | +-------------- Potamilus_fragilis +--+ | +----------- Ptychobranchus +--+ | +-------- Potamilus_purpuratus +--+ | +----- Lampsilis +--+ | +-- Villosa_delumbis ** +--+ +-- Ellipsaria
Krebs et al., 2003 [Publ 2691], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 32 | 10 | 7 | -- |
+----- Ptychobranchus
+--------+
| | +-- Potamilus_fragilis
| +--+
| +-- Medionidus
--+
| +-- Villosa_delumbis **
| +--------+
| | +-- Ellipsaria
+--+
| +-------- Lampsilis_teres
+--+
| +----- Lampsilis_siliquoidea
+--+
| +-- Potamilus_purpuratus
+--+
+-- Potamilus_alatus
