The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)
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Bivalvia | Unionida | Unionoidea | Unionidae | Parreysiinae | Indochinellini genus Scabies Haas, 1911: cladomics 8 Recent valid species (8 total species) Cladomics Summarynumber of species = 8 (8 total) analyzed in 47 cladograms in 29 publications maximum number of terminals in an analysis =
8 (mean = 3.6, median = 3) Basic Monophylynumber of cladograms that test basic monophyly
(i.e., more than 1 included species) = 26 maximum number of included terminals = 8, (mean =
5.69 median = 8) proportion of cladograms that support basic monophyly
= 0.92
Pairwise Cladogram ClusteringThe graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.
Bolotov et al., 2024 [Publ 4369], fig. 2 ** Top Monophyly Cladogram ** ** Top Sister-Group Cladogram ** | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 54 | 21 | 7 | 0 |
+-- Pseudoscabies
+-----------------------------+
| +-- Indonaia
--+
| +----------------------------- Indochinella
| |
+--+ +-------------------------- Radiatula
| |
+--+ +----------------------- Scabiellus
| |
+--+ +-------------------- Harmandia
| |
+--+ +----------------- Unionetta
| |
+--+ +-- Scabies_crispata **
| +-----------+
| | +-- Scabies_mandarinus **
+--+
| +----------- Scabies_nucleus **
| |
+--+ +-- Scabies_humilis **
| +-----+
| | +-- Scabies_pilata **
+--+
| +----- Scabies_phaselus **
+--+
| +-- Scabies_anceps **
+--+
+-- Scabies_scobinatus **
Bolotov et al., 2022 [Publ 4252], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 53 | 19 | 6 | 0 |
+-------------------------------- Indonaia
|
--+ +----------------------------- Indochinella
| |
+--+ +-------------------------- Radiatula
| |
+--+ +----------------------- Scabiellus
| |
+--+ +-------------------- Harmandia
| |
+--+ +----------------- Unionetta
| |
+--+ +-- Scabies_crispata **
| +-----------+
| | +-- Scabies_mandarinus **
+--+
| +----------- Scabies_nucleus **
| |
+--+ +-- Scabies_humilis **
| +-----+
| | +-- Scabies_pilata **
+--+
| +----- Scabies_phaselus **
+--+
| +-- Scabies_anceps **
+--+
+-- Scabies_scobinatus **
Bolotov et al., 2022 [Publ 4239], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 50 | 18 | 6 | 6 |
+-------------------------------------- Indonaia
--+
| +----------------------------------- Indochinella
+--+
| +-------------------------------- Radiatula
+--+
| +----------------------------- Scabiellus
+--+
| +-------------------------- Harmandia
+--+
| +----------------------- Unionetta
+--+
| +-------------------- Scabies_crispata **
+--+
| +----------------- Scabies_mandarinus **
+--+
| +-------------- Scabies_nucleus **
+--+
| +----------- Scabies_pilata **
+--+
| +-------- Scabies_humilis **
+--+
| +----- Scabies_anceps **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_scobinatus **
Pfeiffer et al., 2018 [Publ 3827], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 5 | 8 | 20 | 9 | 6 | 1 |
+-------------------------------- Indonaia
|
--+ +----------------------------- Indochinella
| |
+--+ +-------------------------- Radiatula
| |
+--+ +----------------------- Scabiellus
| |
+--+ +-------------------- Unionetta_fabagina__2
| |
+--+ +----------------- Harmandia
| |
+--+ +-------------- Unionetta_fabagina__1
| |
+--+ +-- Scabies_crispata **
| +--------+
| | +-- Scabies_mandarinus **
+--+
| +-------- Scabies_nucleus **
| |
+--+ +-- Scabies_pilata **
| +--+
| | +-- Scabies_humilis **
+--+
| +-- Scabies_phaselus **
| |
+--+-- Scabies_anceps **
|
+-- Scabies_scobinatus **
Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 622 | 20 | 6 | 10 |
+-------------------- Harmandia
|
--+ +----------------- Unionetta
| |
+--+ +-------------- Scabies_crispata **
| |
| | +----- Scabies_nucleus **
+--+ +-----+
| | | +-- Scabies_mandarinus **
| | +--+
+--+ +-- Scabies_pilata **
|
| +-------- Scabies_humilis **
+--+
| +----- Scabies_anceps **
+--+
| +-- Scabies_scobinatus **
+--+
+-- Scabies_phaselus **
Jeratthitikul et al., 2019 [Publ 3922], fig. 1 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 15 | 10 | 6 | 8 |
+-------------------------------------- Indonaia
--+
| +----------------------------------- Indochinella
+--+
| +-------------------------------- Radiatula
+--+
| +----------------------------- Scabiellus
+--+
| +-------------------------- Harmandia
+--+
| +----------------------- Unionetta
+--+
| +-------------------- Scabies_crispata **
+--+
| +----------------- Scabies_mandarinus **
+--+
| +-------------- Scabies_pilata **
+--+
| +----------- Scabies_nucleus **
+--+
| +-------- Scabies_humilis **
+--+
| +----- Scabies_scobinatus **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_anceps **
Chen et al., 2026 [Publ 4833], fig. 1 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 19 | 6 | 5 | 8 |
+----------------------------- Harmandia
|
--+ +-- Unionetta_xishuangbannaensis
| +-----------------------+
| | +-- Unionetta_fabagina__2
+--+
| +----------------------- Unionetta_fabagina__1
+--+
| +-------------------- Scabies_crispata **
+--+
| +----------------- Scabies_mandarinus **
+--+
| +-------------- Scabies_humilis **
+--+
| +----------- Scabies_pilata **
+--+
| +-------- Scabies_nucleus **
+--+
| +----- Scabies_scobinatus **
+--+
| +-- Scabies_anceps **
+--+
+-- Scabies_phaselus **
Bolotov et al., 2024 [Publ 4369], fig. S1 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 55 | 21 | 7 | 6 |
+-- Pseudoscabies
+-----------------------------------+
| +-- Indonaia
--+
| +----------------------------------- Indochinella
+--+
| +-------------------------------- Radiatula
+--+
| +----------------------------- Scabiellus
+--+
| +-------------------------- Harmandia
+--+
| +----------------------- Unionetta
+--+
| +-------------------- Scabies_crispata **
+--+
| +----------------- Scabies_mandarinus **
+--+
| +-------------- Scabies_nucleus **
+--+
| +----------- Scabies_pilata **
+--+
| +-------- Scabies_humilis **
+--+
| +----- Scabies_anceps **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_scobinatus **
Bolotov et al., 2020 [Publ 4100], fig. 1 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 109 | 13 | 6 | 4 |
+----------------------------- Indonaia
|
--+ +-------------------------- Indochinella
| |
+--+ +----------------------- Radiatula
| |
+--+ +-------------------- Scabiellus
| |
| | +-- Unionetta
+--+ +--------------+
| | +-- Harmandia
| |
+--+ +-------------- Scabies_nucleus **
| |
| | +-- Scabies_crispata **
+--+ +--------+
| | +-- Scabies_mandarinus **
| |
+--+ +-- Scabies_humilis **
| +-----+
| | +-- Scabies_pilata **
+--+
| +----- Scabies_anceps **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_scobinatus **
Bolotov et al., 2022 [Publ 4252], fig. S1 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 53 | 19 | 6 | 10 |
+----------------------------------- Indonaia
|
--+ +-------------------------------- Indochinella
| |
+--+ +----------------------------- Radiatula
| |
+--+ +-------------------------- Scabiellus
| |
+--+ +----------------------- Harmandia
| |
+--+ +-------------------- Unionetta
| |
+--+ +----------------- Scabies_crispata **
| |
+--+ +-- Scabies_humilis **
| +-----------+
| | +-- Scabies_mandarinus **
+--+
| +----------- Scabies_nucleus **
+--+
| +-------- Scabies_pilata **
+--+
| +----- Scabies_anceps **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_scobinatus **
Konopleva et al., 2023 [Publ 4262], fig. 8 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 4 | 8 | 74 | 6 | 6 | 6 |
+----------------------- Indonaia
|
--+ +-------------------- Indochinella
| |
+--+ +----------------- Radiatula
| |
+--+ +-------------- Scabiellus
| |
| | +-- Unionetta
+--+ +--------+
| | +-- Harmandia
| |
| | +-- Scabies_humilis **
+--+ +-----+
| | +-- Scabies_pilata **
| |
| | +----- Scabies_crispata **
| +--+
+--+ | +-- Scabies_nucleus **
| +--+
| +-- Scabies_mandarinus **
|
| +-- Scabies_phaselus **
| |
+-----+-- Scabies_anceps **
|
+-- Scabies_scobinatus **
Bolotov et al., 2022 [Publ 4239], fig. S2 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 50 | 18 | 6 | 10 |
+----------------------------------- Indonaia
|
--+ +-------------------------------- Indochinella
| |
+--+ +----------------------------- Radiatula
| |
+--+ +-------------------------- Scabiellus
| |
+--+ +----------------------- Harmandia
| |
+--+ +-------------------- Unionetta
| |
+--+ +----------------- Scabies_crispata **
| |
+--+ +-- Scabies_humilis **
| +-----------+
| | +-- Scabies_mandarinus **
+--+
| +----------- Scabies_nucleus **
+--+
| +-------- Scabies_pilata **
+--+
| +----- Scabies_anceps **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_scobinatus **
Bolotov et al., 2022 [Publ 4239], fig. S1 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 50 | 18 | 6 | 0 |
+-------------------------------- Indonaia
|
--+ +----------------------------- Indochinella
| |
+--+ +-------------------------- Radiatula
| |
+--+ +----------------------- Scabiellus
| |
+--+ +-------------------- Harmandia
| |
+--+ +----------------- Unionetta
| |
+--+ +-- Scabies_crispata **
| +-----------+
| | +-- Scabies_mandarinus **
+--+
| +----------- Scabies_nucleus **
| |
+--+ +-- Scabies_humilis **
| +-----+
| | +-- Scabies_pilata **
+--+
| +----- Scabies_phaselus **
+--+
| +-- Scabies_anceps **
+--+
+-- Scabies_scobinatus **
Bolotov et al., 2020 [Publ 4100], fig. S1 | iT | tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 5 | 8 | 103 | 12 | 6 | 3 |
+----------------------------------- Indonaia
|
--+ +-------------------------------- Indochinella
| |
| | +----- Radiatula_mouhoti
+--+ +-----------------------+
| | | +-- Radiatula_myitkyinae
| | +--+
+--+ +-- Radiatula_chaudhurii
|
| +-------------------------- Radiatula_crispisulcata
| |
+--+ +----------------------- Scabiellus
| |
+--+ +-------------------- Harmandia
| |
+--+ +----------------- Unionetta
| |
+--+ +-------------- Scabies_mandarinus **
| |
+--+ +----------- Scabies_crispata **
| |
+--+ +-------- Scabies_nucleus **
| |
+--+ +-- Scabies_humilis **
| +--+
| | +-- Scabies_pilata **
+--+
| +-- Scabies_phaselus **
| |
+--+-- Scabies_scobinatus **
|
+-- Scabies_anceps **
Bolotov et al., 2017 [Publ 3761], fig. S1 | iT | tT | bi | riS | oT | roT | roS | iD | | 4 | 7 | 5 | 4 | 135 | 7 | 3 | 9 |
+-------------- Indonaia
|
--+ +-- Indochinella
| +--------+
| | +-- Scabies_crispata **
+--+
| +-------- Radiatula
+--+
| +----- Scabies_mandarinus **
+--+
| +-- Scabies_humilis **
+--+
+-- Scabies_phaselus **
Bolotov et al., 2017 [Publ 3761], fig. S2 | iT | tT | bi | riS | oT | roT | roS | iD | | 4 | 7 | 5 | 4 | 135 | 7 | 3 | 9 |
+-------------- Indonaia
|
--+ +-- Indochinella
| +--------+
| | +-- Scabies_crispata **
+--+
| +-------- Radiatula
+--+
| +----- Scabies_mandarinus **
+--+
| +-- Scabies_humilis **
+--+
+-- Scabies_phaselus **
Ortiz-Sepulveda et al., 2020 [Publ 4101], fig. 1 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 0 | 3 | 33 | 5 | 3 | 6 |
+----------- Indonaia
|
--+ +-------- Indochinella
| |
| | +----- Radiatula_mouhoti
+--+ |
| +----- Radiatula_myitkyinae
| |
+--+----- Radiatula_chaudhurii
|
| +-- Scabies_humilis **
| |
+--+-- Scabies_phaselus **
|
+-- Scabies_mandarinus **
Bolotov et al., 2017 [Publ 3764], fig. S1 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 0 | 3 | 122 | 9 | 3 | 6 |
+-------- Indochinella
|
--+ +----- Radiatula
| |
+--+ +-- Scabies_humilis **
| |
+--+-- Scabies_phaselus **
|
+-- Scabies_mandarinus **
Huang et al., 2019 [Publ 3843], fig. S4 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 1 | 3 | 198 | 9 | 3 | 7 |
+-------------- Indonaia
--+
| +----------- Indochinella
+--+
| +-------- Radiatula
+--+
| +----- Scabies_humilis **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_mandarinus **
Huang et al., 2019 [Publ 3843], fig. S3 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 1 | 3 | 196 | 9 | 3 | 7 |
+-------------- Indonaia
--+
| +----------- Indochinella
+--+
| +-------- Radiatula
+--+
| +----- Scabies_phaselus **
+--+
| +-- Scabies_humilis **
+--+
+-- Scabies_mandarinus **
Konopleva et al., 2019 [Publ 3900], fig. 1 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 0 | 3 | 48 | 9 | 3 | 6 |
+----------- Indonaia
|
--+ +-------- Indochinella
| |
+--+ +----- Radiatula
| |
+--+ +-- Scabies_humilis **
| |
+--+-- Scabies_mandarinus **
|
+-- Scabies_phaselus **
Bolotov et al., 2018 [Publ 3826], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 1 | 3 | 79 | 8 | 3 | 5 |
+-------------- Indonaia
--+
| +----------- Indochinella
+--+
| +-------- Radiatula
+--+
| +----- Scabies_mandarinus **
+--+
| +-- Scabies_humilis **
+--+
+-- Scabies_phaselus **
Ortiz-Sepulveda et al., 2020 [Publ 4101], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 0 | 3 | 33 | 5 | 3 | 6 |
+----- Radiatula_mouhoti
|
+----- Radiatula_myitkyinae
|
+----- Radiatula_chaudhurii
|
--+----- Indochinella
|
+----- Indonaia
|
| +-- Scabies_humilis **
| |
+--+-- Scabies_phaselus **
|
+-- Scabies_mandarinus **
Huang et al., 2019 [Publ 3843], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 3 | 3 | 1 | 3 | 198 | 9 | 3 | 7 |
+-------------- Indonaia
--+
| +----------- Indochinella
+--+
| +-------- Radiatula
+--+
| +----- Scabies_humilis **
+--+
| +-- Scabies_phaselus **
+--+
+-- Scabies_mandarinus **
Dai et al., 2024 [Publ 4386], fig. 5-BI | iT | tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 20 | 0 | 0 | -- |
+-- Scabies_scobinatus **
--+
+-- Scabies_crispata **
Dai et al., 2024 [Publ 4386], fig. 5-ML | iT | tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 20 | 0 | 0 | -- |
+-- Scabies_scobinatus **
--+
+-- Scabies_crispata **
Combosch et al., 2017 [Publ 3806], fig. 1 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 14 | 0 | 0 | -- |
Dai et al., 2024 [Publ 4385], fig. 3-ML | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 0 | 0 | -- |
Wu et al., 2020 [Publ 4121], fig. S1 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 33 | 0 | 0 | -- |
Dai et al., 2026 [Publ 4757], fig. 3 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 33 | 0 | 0 | -- |
Pfeiffer & Graf, 2015 [Publ 3620], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 39 | 2 | 2 | -- |
+----- Indochinella
--+
| +-- Radiatula
+--+
+-- Scabies_mandarinus **
Wu et al., 2021 [Publ 4366], fig. 2_I | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 90 | 4 | 2 | -- |
+----- Indonaia
--+
| +-- Scabies_crispata **
+--+
+-- Radiatula
Xiang et al., 2025 [Publ 4433], fig. 6 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 27 | 0 | 0 | -- |
Sano et al., 2020 [Publ 4138], fig. 3 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 68 | 0 | 0 | -- |
Sano et al., 2020 [Publ 4138], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 68 | 0 | 0 | -- |
Smith, 2021 [Publ 4331], fig. 1 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 19 | 0 | 0 | -- |
Dai et al., 2024 [Publ 4385], fig. 3-BI | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 0 | 0 | -- |
Wu et al., 2020 [Publ 4121], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 33 | 0 | 0 | -- |
Huang et al., 2018 [Publ 3828], fig. 3_5 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 28 | 0 | 0 | -- |
Combosch et al., 2017 [Publ 3806], fig. 2 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 14 | 0 | 0 | -- |
Konopleva et al., 2017 [Publ 3807], fig. 6 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 30 | 2 | 2 | -- |
+----- Indochinella
--+
| +-- Radiatula
+--+
+-- Scabies_mandarinus **
Pfeiffer et al., 2019 [Publ 3937], fig. 6 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 56 | 3 | 3 | -- |
+-------- Indonaia
--+
| +----- Indochinella
+--+
| +-- Scabies_phaselus **
+--+
+-- Harmandia
Sano et al., 2020 [Publ 4138], fig. 4 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 68 | 0 | 0 | -- |
Dai et al., 2024 [Publ 4386], fig. 3 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 13 | 0 | 0 | -- |
Wu et al., 2021 [Publ 4366], fig. 6 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 90 | 4 | 2 | -- |
+----------- Indonaia
--+
| +-------- Radiatula_myitkyinae
+--+
| +----- Scabies_crispata **
+--+
| +-- Radiatula_mouhoti
+--+
+-- Radiatula_chaudhurii
Jeratthitikul et al., 2019 [Publ 3911], fig. 4 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 8 | 0 | 0 | -- |
Pfeiffer et al., 2019 [Publ 3937], fig. 4-5 | iT | tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 56 | 3 | 3 | -- |
+-------- Indonaia
--+
| +----- Indochinella
+--+
| +-- Scabies_phaselus **
+--+
+-- Harmandia
|