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
superfamily HYRIOIDEA Swainson, 1840: cladomics
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 = 93 (124 total)
number of subtaxa (rank: family) = 1
analyzed in 105 cladograms in 55 publications
maximum number of terminals in an analysis = 28 (mean = 4.07, median = 3)
maximum number of subtaxa (family) = 1 (mean = 1, median = 1)
Basic Monophyly
number of cladograms that test basic monophyly (i.e., more than 1 included species) = 61
maximum number of included terminals = 28, (mean = 6.28 median = 5)
maximum number of subtaxa (family) = 1 (mean = 1, median = 1)
proportion of cladograms that support basic monophyly = 1
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
Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14
** Top Monophyly Cladogram ** ** Top Sister-Group Cladogram **
| tT | bi | riS | oT | roT | roS | iD | |
| 28 | 28 | 34 | 1 | 603 | 601 | 2 | 0 |
+----------------------------- Iridininae
|
+-----------------------------------------------------------------+ +-------------------------- Aspathariinae
| | |
| +--+ +----------------------- Etheria
| | |
| +--+ +-------------------- Anodontites_trapesialis
| | |
| +--+ +----------------- Mulleria
| | |
| +--+ +-------------- Lamproscapha
| | |
| | | +----- Monocondylaea
| +--+ +-----+
| | | | +-- Tamsiella_schroeteriana__2
| | | +--+
| +--+ +-- Tamsiella_tamsiana
| |
| | +-------- Mycetopodinae
| | |
| +--+ +-- Anodontites_guanarensis
| | +--+
| | | +-- Fossula
| +--+
| | +-- Anodontites_trigona
| +--+
| +-- Tamsiella_schroeteriana__1
|
| +-------------------------------------------------------------------------------------------- Hyriidae **
| |
| | +-------------------------------- Lamellidens
| | |
| | +-----+ +-- Balwantia
--+ | | | +--------------------------+
| | | | | +-- Trapezidens
| | | +--+
| | | | +-------------------------- Leoparreysiini
| | | | |
| | | +--+ +----------------------- Pseudomulleria
| | | | |
| | | | | +-- Parreysiini
| | | +--+ +-----------------+
| | | | | +-- Coelaturini
| | | | |
| | | +--+ +-- Pseudoscabies
| | | | +--------------+
| | | | | +-- Indonaia
| | | +--+
| | | | +-------------- Indochinella
| | | | |
| | | +--+ +----------- Radiatula
| | +--------------------------------------------------+ | |
| | | | +--+ +-- Songkhlanaia_songkhramensis__2
| | | | | +-----+
| | | | | | +-- Unionetta_fabagina__2
| | | | +--+
| | | | | +----- Harmandia
| | | | +--+
| | | | | +-- Unionetta_fabagina__1
| | | | +--+
| | | | +-- Scabies
| | | |
| | | | +----- Amuranodonta
| | | | +-----------------------------+
| | | | | | +-- Anemina
+--+ | | | +--+
| | | | +-- Buldowskia
| | | |
| | | | +-- Beringiana
| | | | +-----+
| | | | | +-- Pletholophus
| | +--+ +-----------------------+
| | | | | +----- Mimo
| | | | +--+
| | | | | +-- Acudonta
| | | | +--+
| | | | +-- Sinanodonta
| | | |
| | | | +----------------------------- Cristaria
| | +--+ |
| | | | +----------------- Lepidodesmini
| | | | |
| | | | | +----------- Lanceolariini
| | | | | +--+
| | | | +--------+ | | +-------- Pseudoacuticosta_retiaria__2
| | | | | | | +--+
| | | | | | | | +----- Acuticosta_chinensis__2
| | +--+ | | | +--+
| | | | +--+ | +-- Acuticosta_chinensis__1
| | | | | +--+
| | | | | +-- Pseudoacuticosta_retiaria__1
| | | | |
| | | | | +----------- Globunio
| | | | +--+
| | | | | +-------- Aculamprotulina
| | | | +--+
| | +--+ | +----- Middendorffinaiina
| | | +--+
| | | | +-- Unionina
| | | +--+
| | | +-- Oxynaiina
| | |
| | | +-------- Pseudodontoideus
+--+ | +--------------+
| | | | +----- Lasmigona
| | | +--+
| | | | +-- Pterosyna
| | | +--+
| +--+ +-- Arcidens
| |
| | +-------------------- Anodontina
| | |
| | | +-------------- Utterbackia
| | | +--+
| +--+ | | +----------- Pyganodon_grandis
| | | +--+
| | | | +-------- Pyganodon_lacustris
| | | +--+
| | | | +----- Pyganodon_cataracta__2
| +--+ +--+
| | | +-- Pyganodon_cataracta__1
| | +--+
| | +-- Potamilus_fragilis__3
| |
| | +-------------- Utterbackiana
| | |
| +--+ +----------- Simpsonaias
| | |
| | | +-- Alasminota
| +--+ +-----+
| | | +-- Alasmidonta
| | |
| +--+ +-- Prolasmidonta
| | +--+
| | | +-- Platynaias
| +--+
| | +-- Anodontoides
| +--+
| +-- Strophitus
|
| +-------------------------------------------------------------------------------------- Rectidentini
| |
| | +----------------------------------------------------------------------------------- Margaritiferidae
| | |
| | | +-------------------- Chamberlainiini
| | | |
| | | +--+ +----------------- Parvasolenaia
| | | | | |
| | | | +--+ +-------------- Pseudopostolata
| | | | | |
| | | | +--+ +----------- Koreosolenaia
| | | | | |
| | | | +--+ +-- Cosmopseudodon
| | | | | +-----+
| | | | | | +-- Guiunio
+--+ | | +--+
| | +--------+ | +----- Ptychorhynchus
| | | | +--+
| | | | | +-- Obovalis
| | | | +--+
| | | | +-- Postolata
| | | |
| | | | +-- Caudiculatus
| | | | +--------------+
| | | | | +-- Lamprotulini
| | | | |
| | | +-----+ +-------------- Sinosolenaia_carinata
| | | | |
| | | | | +----- Gonidea
| | | +--+ +-----+
+--+ +-----------------------------------------------+ | | | +-- Microcondylaea
| | | | | +--+
| | | +--+ +-- Leguminaia
| | | |
| | | | +-------- Inversidens
| | | +--+
| | | | +----- Sinosolenaia_recognita
| | | +--+
| | | | +-- Sinosolenaia_iridinea
| | | +--+
| | | +-- Sinosolenaia_oleivora
| | |
| | | +----------------------------- Indopseudodontina
| | | |
| | | | +-- Lannanaia
| | +--+ +-----------------------+
| | | | +-- Pilsbryoconcha
| | | |
| | | | +----- Songkhlanaia_tamodienica
| | +--+ +-----------------+
| | | | | +-- Songkhlanaia_moreleti
| | | | +--+
| | | | +-- Songkhlanaia_songkhramensis__1
| | | |
| | +--+ +-- Nyeinchanconcha
| | | +-----------------+
| | | | +-- Sundadontina
| | | |
+--+ | | +-- Isannaia
| +--+ +--+
| | | +-- Namkongnaia
| | +-----------+
| | | | +-- Thaiconcha
| | | +--+
| +--+ +-- Bineurus
| |
| | +-------------- Pseudodon_walpolei
| | |
| +--+ +----------- Pseudodon_vagulus
| | |
| | | +----- Pseudodon_vondembuschianus
| +--+ +--+
| | | | +-- Pseudodon_mekongi
| | | +--+
| +--+ +-- Schepmaniini
| |
| | +----- Pseudodon_lenyanensis__2
| +--+
| | +-- Pseudodon_inoscularis
| +--+
| +-- Pseudodon_lenyanensis__1
|
| +-- Quadrulini
| +--------------------------------------------------------------------------+
| | +-- Pleurobemini
| |
| | +----- 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
Santos-Neto et al., 2016 [Publ 3739], fig. 2a_b
| tT | bi | riS | oT | roT | roS | iD | |
| 19 | 19 | 20 | 1 | 1 | 0 | 0 | 22.5 |
1
Santos-Neto et al., 2016 [Publ 3739], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 19 | 19 | 21 | 1 | 1 | 0 | 0 | 25.5 |
1
Huang et al., 2019 [Publ 3843], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 14 | 14 | 12 | 1 | 186 | 184 | 2 | 39 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Huang et al., 2019 [Publ 3843], fig. S3
| tT | bi | riS | oT | roT | roS | iD | |
| 14 | 14 | 12 | 1 | 184 | 182 | 2 | 41 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Huang et al., 2019 [Publ 3843], fig. S4
| tT | bi | riS | oT | roT | roS | iD | |
| 14 | 14 | 9 | 1 | 186 | 184 | 2 | 38 |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Graf et al., 2015 [Publ 3619], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 14 | 14 | 12 | 1 | 6 | 5 | 2 | 37 |
+----- Hyriidae ** --+ | +-- Unionoidea +--+ +-- Etherioidea
Walker et al., 2006 [Publ 2666], fig. 6
| tT | bi | riS | oT | roT | roS | iD | |
| 10 | 10 | 7 | 1 | 26 | 25 | 2 | 29 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Marshall et al., 2014 [Publ 3593], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 10 | 10 | 8 | 1 | 4 | 4 | 1 | 31.5 |
+-- Margaritiferidae
+--+
| +-- Unioninae
--+
| +-- Ambleminae
+--+
+-- Hyriidae **
Bolotov et al., 2017 [Publ 3761], fig. S2
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 129 | 127 | 2 | 38 |
+-- Chamberlainia
+--------------------+
| +-- Margaritiferidae
|
| +-- Etherioidea
--+--------------------+
| +-- Hyriidae **
|
| +-------------------- Parreysiinae
+--+
| +----------------- Unioninae
+--+
| +-------------- Rectidentini
+--+
| +----------- Ambleminae
+--+
| +-------- Pseudodontini
+--+
| +----- Gonideini
+--+
| +-- Sinohyriopsis
+--+
+-- Lamprotulini
Bolotov et al., 2016 [Publ 3743], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 21 | 18 | 2 | 38 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Hoeh et al., 2009 [Publ 2716], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 38 | 37 | 2 | 32.5 |
+-------------------------- Hyriidae ** | | +-------- Iridinidae --+ +--------------+ | | | +----- Etheria | | +--+ | | | +-- Mulleria +--+ +--+ | +-- Mycetopodidae | | +-------------------- Margaritiferidae +--+ | +----------------- Parreysiinae +--+ | +-------------- Pseudomulleria +--+ | +----------- Unioninae +--+ | +-------- Rectidentini +--+ | +----- Ambleminae +--+ | +-- Gonideini +--+ +-- Pseudodontini
Hoeh et al., 2009 [Publ 2716], fig. 5
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 37 | 36 | 2 | 32.5 |
+----------------- Parreysiinae | | +----- Margaritiferidae --+ +--------+ | | | +-- Hyriidae ** | | +--+ +--+ +-- Etherioidea | | +----------- Pseudodontini +--+ | +-------- Rectidentini +--+ | +----- Unioninae +--+ | +-- Gonideini +--+ +-- Ambleminae
Graf & Cummings, 2006 [Publ 1968], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 38 | 37 | 2 | 32.5 |
+----- Unionoidea --+ | +-- Hyriidae ** +--+ +-- Etherioidea
Bolotov et al., 2016 [Publ 3743], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 21 | 18 | 2 | 38 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Bolotov et al., 2017 [Publ 3761], fig. S1
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 129 | 127 | 2 | 38 |
+-------- Margaritiferidae --+ | +----- Unionidae +--+ | +-- Etherioidea +--+ +-- Hyriidae **
Hoeh et al., 2009 [Publ 2716], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 9 | 9 | 7 | 1 | 37 | 36 | 2 | 32.5 |
+-------------------------- Hyriidae ** | | +-- Etheria | +--+ --+ | +-- Mulleria | +-----------------+ | | | +-- Mycetopodidae | | +--+ +--+ +-- Iridinidae | | +-------------------- Parreysiinae +--+ | +----------------- Pseudomulleria +--+ | +-------------- Margaritiferidae +--+ | +----------- Pseudodontini +--+ | +-------- Rectidentini +--+ | +----- Unioninae +--+ | +-- Gonideini +--+ +-- Ambleminae
Baker et al., 2004 [Publ 2685], fig. 2-3
| tT | bi | riS | oT | roT | roS | iD | |
| 8 | 8 | 9 | 1 | 1 | 1 | 1 | 35 |
+-- Unionoidea --+ +-- Hyriidae **
Santos-Neto et al., 2016 [Publ 3739], fig. 3_5
| tT | bi | riS | oT | roT | roS | iD | |
| 7 | 7 | 6 | 1 | 0 | 0 | 0 | 29 |
1
Baker et al., 2003 [Publ 2690], fig. 5
| tT | bi | riS | oT | roT | roS | iD | |
| 7 | 7 | 5 | 1 | 1 | 1 | 1 | 37 |
+-- Unionoidea --+ +-- Hyriidae **
Graf & Ó Foighil, 2000 [Publ 2649], fig. 1_2
| tT | bi | riS | oT | roT | roS | iD | |
| 6 | 6 | 5 | 1 | 5 | 5 | 1 | 36.5 |
+-- Margaritiferidae | --+-- Unionidae | +-- Hyriidae **
Campbell & Lydeard, 2026 [Publ 4810], fig. 3_5
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 136 | 135 | 2 | 33 |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Guerra et al., 2019 [Publ 4168], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 36 | 35 | 2 | 33 |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Roe & Hoeh, 2003 [Publ 2655], fig. 4f2
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 2 | 1 | 26 | 25 | 2 | 36 |
+----- Unionoidea --+ | +-- Etherioidea +--+ +-- Hyriidae **
Roe & Hoeh, 2003 [Publ 2655], fig. 4f3
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 26 | 25 | 2 | 33 |
+-------- Margaritiferidae --+ | +----- Unionidae +--+ | +-- Etherioidea +--+ +-- Hyriidae **
Hoeh et al., 2001 [Publ 2652], fig. 14f4
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 26 | 25 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Hoeh et al., 2001 [Publ 2652], fig. 14f2
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 1 | 1 | 26 | 25 | 2 | 35 |
+----------------- Margaritiferidae | | +-------------- Amblemini | | | +-------------- Parreysiinae | | | +-------------- Eurynia --+ | | +-------------- Fusconaia | | | +-------------- Gonideinae | | | +-------------- Pleurobema | | +--+-------------- Quadrulini | | +----------- Ortmanniana +--+ | | +-------- Glebula | +--+ | | +----- Potamilus | +--+ | | +-- Ligumia | +--+ | +-- Toxolasma | | +----------- Cyrtonaias +--+ | +-------- Unionini +--+ | +----- Anodontini +--+ | +-- Hyriidae ** +--+ +-- Etherioidea
Guerra et al., 2019 [Publ 4168], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 36 | 35 | 2 | 35 |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Hoeh et al., 2002 [Publ 2653], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 26 | 25 | 2 | 33.5 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Santos-Neto et al., 2018 [Publ 3888], fig. 5
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 0 | 0 | 0 | 31 |
1
Bogan & Hoeh, 2000 [Publ 2650], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 5 | 5 | 3 | 1 | 29 | 28 | 2 | 33 |
+-------------------- Hyriidae ** | | +-------- Iridinidae --+ +--------+ | | | +----- Etheria | | +--+ | | | +-- Mulleria +--+ +--+ | +-- Mycetopodidae | | +-------------- Parreysiinae | | +--+ +----------- Pseudomulleria | | +--+ +-------- Anodontini | | +--+ +----- Gonideinae | | +--+----- Unionini | | +-- Margaritiferidae +--+ +-- Ambleminae
Whelan et al., 2011 [Publ 3380], fig. 1-ML
| tT | bi | riS | oT | roT | roS | iD | |
| 4 | 4 | 2 | 1 | 32 | 31 | 2 | 34 |
+-------- Hyriidae ** --+ | +----- Margaritiferidae +--+ | +-- Etherioidea +--+ +-- Unionidae
Graf, 2002 [Publ 2651], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 4 | 4 | 2 | 1 | 14 | 13 | 1 | 31.5 |
+-- Hyriidae ** --+ +-- Unionoidea
Whelan et al., 2011 [Publ 3380], fig. 1-MP
| tT | bi | riS | oT | roT | roS | iD | |
| 4 | 4 | 2 | 1 | 32 | 31 | 2 | 34 |
+-------- Etherioidea --+ | +----- Unionidae +--+ | +-- Margaritiferidae +--+ +-- Hyriidae **
Klunzinger et al., 2022 [Publ 4259], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 4 | 4 | 2 | 1 | 0 | 0 | 0 | 34 |
1
Pfeiffer & Graf, 2015 [Publ 3620], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 4 | 4 | 2 | 1 | 36 | 35 | 2 | 34 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Graf, 2000 [Publ 2647], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 4 | 4 | 2 | 1 | 13 | 12 | 2 | 34.5 |
+-------------- Margaritiferidae | | +-------- Unionini --+ +--+ | | | +----- Pleurobemini | | +--+ | | | +-- Anodontini +--+ +--+ | +-- Lampsilini | | +-------- Parreysiini +--+ | +----- Coelaturini +--+ | +-- Hyriidae ** +--+ +-- Etherioidea
Klunzinger et al., 2021 [Publ 4086], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 4 | 4 | 2 | 1 | 2 | 2 | 1 | 32 |
+-- Unionoidea --+ +-- Hyriidae **
Sano et al., 2020 [Publ 4138], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 65 | 63 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Sonowal et al., 2021 [Publ 4221], fig. 3a
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 0 | 1 | 12 | 9 | 1 | 34 |
1
Wu et al., 2020 [Publ 4121], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 31 | 29 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Wu et al., 2020 [Publ 4121], fig. S1
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 31 | 29 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Sano et al., 2020 [Publ 4138], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 65 | 63 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Sonowal et al., 2021 [Publ 4221], fig. 3b
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 0 | 1 | 12 | 9 | 1 | 34 |
+-- Hyriidae ** --+ +-- Unionoidea
Lopes-Lima et al., 2018 [Publ 3829], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 25 | 23 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Wu et al., 2021 [Publ 4366], fig. 6
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 88 | 86 | 2 | 35 |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Combosch et al., 2017 [Publ 3806], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 12 | 10 | 2 | 33 |
+-------- Hyriidae ** --+ | +----- Unionidae +--+ | +-- Margaritiferidae +--+ +-- Etherioidea
Huang et al., 2018 [Publ 3828], fig. 3_5
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 26 | 24 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Sano et al., 2020 [Publ 4138], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 65 | 63 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Olivera-Hyde et al., 2020 [Publ 4249], fig. 10
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 0 | 0 | 0 | 33 |
1
Konopleva et al., 2017 [Publ 3807], fig. 6
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 28 | 26 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Lopes-Lima et al., 2018 [Publ 3829], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 25 | 23 | 2 | 33 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Sonowal et al., 2021 [Publ 4221], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 13 | 10 | 1 | 33 |
+-- Hyriidae ** --+ +-- Unionoidea
Combosch et al., 2017 [Publ 3806], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 12 | 10 | 2 | 35 |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Wu et al., 2021 [Publ 4366], fig. 2_I
| tT | bi | riS | oT | roT | roS | iD | |
| 3 | 3 | 1 | 1 | 88 | 86 | 2 | 35 |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Guerra et al., 2019 [Publ 4168], fig. 1-M
| tT | bi | riS | oT | roT | roS | iD | |
| 2 | 2 | 0 | 1 | 20 | 20 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Lopes-Lima et al., 2020 [Publ 4080], fig. 2-4
| tT | bi | riS | oT | roT | roS | iD | |
| 2 | 2 | 0 | 1 | 120 | 118 | 2 | -- |
+-------- Hyriidae ** --+ | +----- Margaritiferidae +--+ | +-- Etherioidea +--+ +-- Unionidae
Guerra et al., 2019 [Publ 4168], fig. 2-M
| tT | bi | riS | oT | roT | roS | iD | |
| 2 | 2 | 0 | 1 | 20 | 20 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Campbell & Lydeard, 2026 [Publ 4810], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 2 | 2 | 0 | 1 | 45 | 44 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Lopes-Lima et al., 2017 [Publ 3745], fig. 1-2
| tT | bi | riS | oT | roT | roS | iD | |
| 2 | 2 | 0 | 1 | 78 | 77 | 2 | -- |
+-------- Hyriidae ** --+ | +----- Margaritiferidae +--+ | +-- Etherioidea +--+ +-- Unionidae
Chen et al., 2025 [Publ 4455], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 2 | 2 | 0 | 1 | 24 | 22 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Wu et al., 2020 [Publ 4121], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 48 | 47 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Plazzi et al., 2011 [Publ 3676], fig. 5
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 17 | 17 | 1 | -- |
+----------- Cristariina | | +-------- Lanceolariini | | --+ +-------- Anodonta | | | +-------- Unionini | | +--+ +-- Alasmidontina +-----+ | +-- Pseudanodonta | | +----- Ambleminae | | +--+ +-- Hyriidae ** | | +--+-- Lamprotulini | +-- Chamberlainiini
Bieler et al., 2014 [Publ 3622], fig. 28
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 4 | 3 | 2 | -- |
+-- Etherioidea
+--+
| +-- Margaritiferidae
--+
| +-- Unionidae
+--+
+-- Hyriidae **
Wu et al., 2023 [Publ 4339], fig. 4_S5-aa
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 95 | 94 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Graf & Ó Foighil, 2000 [Publ 2648], fig. 2B
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 29 | 28 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Wu et al., 2021 [Publ 4366], fig. 3-BI
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 73 | 72 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Wu et al., 2024 [Publ 4350], fig. 8b_S2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 91 | 90 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Wu et al., 2019 [Publ 3864], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 45 | 45 | 2 | -- |
+-------- Unionidae --+ | +----- Margaritiferidae +--+ | +-- Hyriidae ** +--+ +-- Etherioidea
González & Giribet, 2015 [Publ 3632], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 5 | 3 | 2 | -- |
+-- Etherioidea
+--+
| +-- Margaritiferidae
--+
| +-- Unionidae
+--+
+-- Hyriidae **
Wu et al., 2022 [Publ 4238], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 73 | 73 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Bieler et al., 2014 [Publ 3622], fig. 26_27_29_30
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 4 | 3 | 2 | -- |
+-------- Hyriidae ** --+ | +----- Unionidae +--+ | +-- Etherioidea +--+ +-- Margaritiferidae
González & Giribet, 2015 [Publ 3632], fig. 6
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 5 | 3 | 2 | -- |
+-------- Hyriidae ** --+ | +----- Unionidae +--+ | +-- Etherioidea +--+ +-- Margaritiferidae
González & Giribet, 2015 [Publ 3632], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 5 | 3 | 2 | -- |
+----- Unionidae | --+----- Hyriidae ** | | +-- Etherioidea +--+ +-- Margaritiferidae
Guerra et al., 2017 [Publ 3772], fig. 2b-M
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 7 | 7 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Huang et al., 2019 [Publ 3843], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 55 | 54 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Guerra et al., 2017 [Publ 3772], fig. 2a-M
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 7 | 7 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Walker et al., 2006 [Publ 2666], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 14 | 13 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Chase et al., 2018 [Publ 3847], fig. 5
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 9 | 9 | 1 | -- |
+----- Unionidae --+ | +-- Margaritiferidae +--+ +-- Hyriidae **
Wu et al., 2023 [Publ 4339], fig. 4_S5-nt
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 95 | 94 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Chase et al., 2018 [Publ 3861], fig. 1
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 10 | 10 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Guerra et al., 2017 [Publ 3772], fig. 2b
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 24 | 23 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Hoeh et al., 2002 [Publ 2673], fig. 1-M
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 8 | 8 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Huang et al., 2018 [Publ 3828], fig. 6
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 41 | 40 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Wu et al., 2021 [Publ 4366], fig. 3-ML
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 73 | 72 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Wu et al., 2024 [Publ 4350], fig. 8a_S1
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 91 | 90 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Pfeiffer et al., 2019 [Publ 3937], fig. 6
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 56 | 55 | 2 | -- |
+-------- Etherioidea --+ | +----- Unionidae +--+ | +-- Hyriidae ** +--+ +-- Margaritiferidae
Chase et al., 2018 [Publ 3847], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 38 | 38 | 1 | -- |
+-------------- Margaritiferidae | --+ +----------- Unioninae | | +--+ +-------- Lampsilini | | +--+ +-- Hyriidae ** | +--+ | | +-- Amblemini +--+ | +-- Pleurobemini +--+ +-- Quadrulini
Pfeiffer & Graf, 2013 [Publ 3531], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 30 | 30 | 1 | -- |
+-- Margaritiferidae | --+-- Hyriidae ** | +-- Unionidae
Walker et al., 2006 [Publ 2666], fig. 6-M
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 13 | 13 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Chase et al., 2018 [Publ 3861], fig. 1-M
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 10 | 10 | 1 | -- |
+----- Unionidae --+ | +-- Margaritiferidae +--+ +-- Hyriidae **
Wu et al., 2021 [Publ 4366], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 68 | 67 | 2 | -- |
+----- Hyriidae ** --+ | +-- Etherioidea +--+ +-- Unionoidea
Elderkin et al., 2016 [Publ 3693], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 2 | 2 | 1 | -- |
+-- Etheriidae | --+-- Iridinidae | +-- Hyriidae **
Dai et al., 2025 [Publ 4440], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 87 | 87 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Walker et al., 2006 [Publ 2666], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 14 | 13 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Hoeh et al., 2002 [Publ 2673], fig. 1-F
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 8 | 8 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
González & Giribet, 2015 [Publ 3632], fig. 4
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 5 | 3 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Breton et al., 2011 [Publ 3376], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 37 | 37 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Chen et al., 2025 [Publ 4448], fig. 3
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 85 | 85 | 2 | -- |
+-------- Etherioidea --+ | +----- Unionidae +--+ | +-- Hyriidae ** +--+ +-- Margaritiferidae
Guerra et al., 2017 [Publ 3772], fig. 2a
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 24 | 23 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Graf & Ó Foighil, 2000 [Publ 2648], fig. 2A
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 29 | 28 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Huang et al., 2019 [Publ 3843], fig. 3-M
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 22 | 22 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Walker et al., 2006 [Publ 2666], fig. 2
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 14 | 13 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
Pfeiffer et al., 2019 [Publ 3937], fig. 4-5
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 56 | 55 | 2 | -- |
+----- Etherioidea --+ | +-- Hyriidae ** +--+ +-- Unionoidea
Huang et al., 2018 [Publ 3828], fig. 6-M
| tT | bi | riS | oT | roT | roS | iD | |
| 1 | 1 | 0 | 1 | 16 | 16 | 1 | -- |
+-- Hyriidae ** --+ +-- Unionoidea
