The Freshwater Mussels (Unionoida) of the World (and other less consequential bivalves)
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Bivalvia | Unionida | Unionoidea | Unionidae | Ambleminae | Quadrulini | Quadrulina genus Pustulosa Frierson, 1927: cladomics
8 Recent valid species (9 total species) Cladomics Summarynumber of species = 8 (9 total)
analyzed in 42 cladograms in 24 publications
maximum number of terminals in an analysis =
8 (mean = 3.14, median = 2) Basic Monophylynumber of cladograms that test basic monophyly
(i.e., more than 1 included species) = 25
maximum number of included terminals = 8, (mean =
4.6 median = 4)
proportion of cladograms that support basic monophyly
= 0.24
Pairwise Cladogram ClusteringThe graphs below depict in two dimensions all pair-wise ingroup distances within the cladome. See the Cladmoics FARQ for more information.
Lopes-Lima et al., 2019 [Publ 3901], fig. 3 ** Top Monophyly Cladogram ** ** Top Sister-Group Cladogram ** iT| tT | bi | riS | oT | roT | roS | iD | | 8 | 9 | 7 | 8 | 21 | 9 | 3 | 0 | +----------------------- Theliderma
|
--+ +-------------------- Quadrula
| |
+--+ +----------------- Pustulosa_kieneriana **
| |
| | +-- Pustulosa_infucata **
+--+ +-----------+
| | +-- Pustulosa_kleiniana **
| |
+--+ +-- Pustulosa_pustulosa **
| +--------+
| | +-- Pustulosa_succissa **
+--+
| +-------- Cyclonaias
+--+
| +----- Pustulosa_necki **
+--+
| +-- Pustulosa_petrina **
+--+
+-- Pustulosa_nodulata **
Johnson et al., 2018 [Publ 3844], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 8 | 9 | 7 | 8 | 8 | 4 | 3 | 2 | +-------------------- Theliderma
|
--+ +----------------- Quadrula
| |
| | +----- Pustulosa_kieneriana **
+--+ +--------+
| | | +-- Pustulosa_infucata **
| | +--+
| | +-- Pustulosa_kleiniana **
+--+
| +-- Pustulosa_pustulosa **
| +--------+
| | +-- Pustulosa_succissa **
+--+
| +-------- Cyclonaias
+--+
| +----- Pustulosa_necki **
+--+
| +-- Pustulosa_nodulata **
+--+
+-- Pustulosa_petrina **
Campbell & Lydeard, 2026 [Publ 4810], fig. 6-14 iT| tT | bi | riS | oT | roT | roS | iD | | 8 | 9 | 7 | 8 | 624 | 8 | 3 | 6 | +-- Theliderma_johnsoni
+--------+
| +-- Theliderma_metanevra
|
+-----+ +-- Quadrula_nobilis
| | +-----+
| | | +-- Quadrula_verrucosa
| +--+
| | +----- Theliderma_cylindrica
| +--+
| | +-- Quadrula_fragosa
--+ +--+
| +-- Quadrula_quadrula
|
| +----- Pustulosa_kieneriana **
| +--------+
| | | +-- Pustulosa_infucata **
| | +--+
+--+ +-- Pustulosa_kleiniana **
|
| +----------- Cyclonaias
| |
+--+ +-- Pustulosa_pustulosa **
| +-----+
| | +-- Pustulosa_succissa **
+--+
| +----- Pustulosa_nodulata **
+--+
| +-- Pustulosa_necki **
+--+
+-- Pustulosa_petrina **
Lopes-Lima et al., 2019 [Publ 3901], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 8 | 9 | 7 | 8 | 20 | 6 | 3 | 6 | +----------------------- Quadrula_quadrula
|
| +-- Quadrula_verrucosa
--+ +-----------------+
| | +-- Quadrula_nobilis
| |
+--+ +----------------- Theliderma
| |
| | +----- Pustulosa_kieneriana **
+--+ +--------+
| | | +-- Pustulosa_infucata **
| | +--+
+--+ +-- Pustulosa_kleiniana **
|
| +----------- Cyclonaias
| |
+--+ +-- Pustulosa_pustulosa **
| +-----+
| | +-- Pustulosa_succissa **
+--+
| +----- Pustulosa_nodulata **
+--+
| +-- Pustulosa_necki **
+--+
+-- Pustulosa_petrina **
Lopes-Lima et al., 2019 [Publ 3901], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 8 | 9 | 7 | 8 | 18 | 6 | 3 | 2 | +----------------------- Theliderma
|
--+ +-------------------- Quadrula_quadrula
| |
| | +-- Quadrula_verrucosa
+--+ +--------------+
| | +-- Quadrula_nobilis
| |
| | +----- Pustulosa_kieneriana **
+--+ +--------+
| | | +-- Pustulosa_infucata **
| | +--+
| | +-- Pustulosa_kleiniana **
+--+
| +-- Pustulosa_pustulosa **
| +--------+
| | +-- Pustulosa_succissa **
+--+
| +-------- Cyclonaias
+--+
| +----- Pustulosa_necki **
+--+
| +-- Pustulosa_petrina **
+--+
+-- Pustulosa_nodulata **
Neemuchwala et al., 2023 [Publ 4292], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 8 | 8 | 6 | 8 | 19 | 9 | 3 | 9 | +-------------------- Theliderma
|
--+ +----------------- Cyclonaias
| |
+--+ +-------------- Quadrula
| |
| | +----- Pustulosa_nodulata **
+--+ +-----+
| | | +-- Pustulosa_necki **
| | +--+
| | +-- Pustulosa_petrina **
+--+
| +-- Pustulosa_kieneriana **
| +-----+
| | +-- Pustulosa_pustulosa **
+--+
| +----- Pustulosa_succissa **
+--+
| +-- Pustulosa_infucata **
+--+
+-- Pustulosa_kleiniana **
Serb et al., 2003 [Publ 2659], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 6 | 6 | 3 | 6 | 23 | 7 | 2 | 4 | +-------------- Theliderma
|
--+ +----------- Quadrula
| |
+--+ +-------- Pustulosa_kieneriana **
| |
| +-------- Pustulosa_infucata **
+--+
| +-- Pustulosa_pustulosa **
| +--+
| | +-- Pustulosa_succissa **
+--+
| +-- Pustulosa_nodulata **
+--+
+-- Pustulosa_petrina **
Szumowski et al., 2012 [Publ 3569], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 6 | 6 | 4 | 6 | 8 | 7 | 2 | 3 | +-------------- Quadrula
|
--+ +----------- Pustulosa_kieneriana **
| |
+--+ +-------- Pustulosa_infucata **
| |
+--+ +-- Pustulosa_pustulosa **
| +--+
| | +-- Pustulosa_succissa **
+--+
| +-- Pustulosa_nodulata **
+--+
+-- Pustulosa_petrina **
Serb et al., 2003 [Publ 2659], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 6 | 6 | 4 | 6 | 23 | 7 | 2 | 3 | +-- Quadrula_verrucosa
+-----------------+
| +-- Quadrula_nobilis
--+
| +----------------- Quadrula_quadrula
| |
+--+ +-------------- Theliderma
| |
+--+ +----------- Pustulosa_kieneriana **
| |
+--+ +-------- Pustulosa_infucata **
| |
+--+ +-- Pustulosa_succissa **
| +--+
| | +-- Pustulosa_pustulosa **
+--+
| +-- Pustulosa_nodulata **
+--+
+-- Pustulosa_petrina **
Campbell & Lydeard, 2012 [Publ 3308], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 6 | 4 | 4 | 39 | 2 | 2 | 5 | +-- Pustulosa_infucata **
+--------+
| +-- Pustulosa_kleiniana **
--+
| +-------- Quadrula
+--+
| +----- Cyclonaias
+--+
| +-- Pustulosa_succissa **
+--+
+-- Pustulosa_pustulosa **
Campbell & Lydeard, 2012 [Publ 3308], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 5 | 3 | 4 | 38 | 2 | 2 | 4 | +----------- Quadrula
|
--+ +-- Pustulosa_infucata **
| +-----+
| | +-- Pustulosa_kleiniana **
+--+
| +----- Cyclonaias
+--+
| +-- Pustulosa_succissa **
+--+
+-- Pustulosa_pustulosa **
Campbell et al., 2005 [Publ 2660], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 5 | 2 | 4 | 98 | 5 | 3 | 5 | +-------------- Theliderma
|
--+ +----------- Quadrula_quadrula
| |
| | +-- Quadrula_nobilis
+--+ +-----+
| | +-- Quadrula_verrucosa
| |
+--+ +----- Pustulosa_kieneriana **
| |
| | +-- Cyclonaias
+--+--+
| +-- Pustulosa_succissa **
|
| +-- Pustulosa_infucata **
+--+
+-- Pustulosa_kleiniana **
Campbell & Lydeard, 2012 [Publ 3309], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 5 | 3 | 4 | 69 | 5 | 3 | 4 | +-- Quadrula_verrucosa
+-----------+
| +-- Quadrula_nobilis
|
--+ +----------- Quadrula_quadrula
| |
| +----------- Theliderma
+--+
| +-- Pustulosa_infucata **
| +-----+
| | +-- Pustulosa_kleiniana **
+--+
| +----- Cyclonaias
+--+
| +-- Pustulosa_pustulosa **
+--+
+-- Pustulosa_succissa **
Campbell & Lydeard, 2012 [Publ 3308], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 5 | 3 | 4 | 38 | 2 | 2 | 4 | +----------- Quadrula
|
--+ +-- Pustulosa_infucata **
| +-----+
| | +-- Pustulosa_kleiniana **
+--+
| +----- Cyclonaias
+--+
| +-- Pustulosa_succissa **
+--+
+-- Pustulosa_pustulosa **
Campbell & Lydeard, 2012 [Publ 3309], fig. 1-2 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 5 | 3 | 4 | 69 | 5 | 3 | 4 | +----------------- Theliderma
|
--+ +-------------- Quadrula_verrucosa
| |
| | +-- Quadrula_quadrula
+--+ +--------+
| | +-- Quadrula_nobilis
| |
+--+ +-- Pustulosa_infucata **
| +-----+
| | +-- Pustulosa_kleiniana **
+--+
| +----- Cyclonaias
+--+
| +-- Pustulosa_pustulosa **
+--+
+-- Pustulosa_succissa **
Campbell & Lydeard, 2012 [Publ 3308], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 5 | 3 | 4 | 38 | 2 | 2 | 4 | +----------- Quadrula
|
--+ +-- Pustulosa_infucata **
| +-----+
| | +-- Pustulosa_kleiniana **
+--+
| +----- Cyclonaias
+--+
| +-- Pustulosa_succissa **
+--+
+-- Pustulosa_pustulosa **
Campbell et al., 2005 [Publ 2660], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 4 | 5 | 2 | 4 | 98 | 5 | 3 | 5 | +----------- Theliderma
|
--+ +-------- Quadrula
| |
+--+ +----- Pustulosa_kieneriana **
| |
| | +-- Cyclonaias
+--+--+
| +-- Pustulosa_succissa **
|
| +-- Pustulosa_infucata **
+--+
+-- Pustulosa_kleiniana **
Zhou et al., 2007 [Publ 2658], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 3 | 4 | 2 | 3 | 15 | 4 | 2 | 9 | +-------------- Theliderma
--+
| +----------- Quadrula_quadrula
+--+
| +-------- Pustulosa_infucata **
+--+
| +----- Quadrula_verrucosa
+--+
| +-- Pustulosa_kieneriana **
+--+
+-- Pustulosa_succissa **
Boyer et al., 2011 [Publ 2718], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 3 | 0 | 2 | 33 | 4 | 3 | 6 | +-------------- Quadrula_quadrula
--+
| +----------- Quadrula_verrucosa
+--+
| +-------- Theliderma
+--+
| +----- Pustulosa_pustulosa **
+--+
| +-- Cyclonaias
+--+
+-- Pustulosa_nodulata **
Boyer et al., 2011 [Publ 2718], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 3 | 0 | 2 | 36 | 4 | 3 | 6 | +----------- Theliderma
--+
| +-------- Quadrula
+--+
| +----- Pustulosa_pustulosa **
+--+
| +-- Cyclonaias
+--+
+-- Pustulosa_nodulata **
Lydeard et al., 2000 [Publ 2654], fig. 2a iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 12 | 1 | 1 | -- | +----- Quadrula
--+
| +-- Pustulosa_infucata **
+--+
+-- Pustulosa_succissa **
Lydeard et al., 2000 [Publ 2654], fig. 2b iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 3 | 0 | 2 | 12 | 1 | 1 | 8 | +-- Quadrula
|
--+-- Pustulosa_infucata **
|
+-- Pustulosa_succissa **
Lydeard et al., 2000 [Publ 2654], fig. 2d iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 3 | 0 | 2 | 12 | 1 | 1 | 8 | +-- Quadrula
|
--+-- Pustulosa_infucata **
|
+-- Pustulosa_succissa **
Pfeiffer et al., 2016 [Publ 3660], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 3 | 0 | 2 | 36 | 3 | 2 | 6 | +----------- Quadrula_quadrula
--+
| +-------- Quadrula_verrucosa
+--+
| +----- Pustulosa_pustulosa **
+--+
| +-- Pustulosa_petrina **
+--+
+-- Cyclonaias
Lydeard et al., 2000 [Publ 2654], fig. 2c iT| tT | bi | riS | oT | roT | roS | iD | | 2 | 2 | 0 | 2 | 12 | 1 | 1 | -- | +----- Quadrula
--+
| +-- Pustulosa_infucata **
+--+
+-- Pustulosa_succissa **
Freeman et al., 2025 [Publ 4443], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 11 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Inoue et al., 2020 [Publ 3941], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 41 | 1 | 1 | -- | +-- Pustulosa_petrina **
--+
+-- Quadrula
Freeman et al., 2025 [Publ 4443], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 11 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Inoue et al., 2020 [Publ 3941], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 40 | 1 | 1 | -- | +-- Pustulosa_petrina **
--+
+-- Quadrula
Porto-Hannes et al., 2021 [Publ 4227], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 10 | 1 | 1 | -- | -- Pustulosa_pustulosa **
Inoue et al., 2018 [Publ 3858], fig. 1_3a-c iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 5 | 0 | 2 | 104 | 4 | 3 | 11 | +-- Cyclonaias
|
+-- Pustulosa_pustulosa__1 **
|
+-- Theliderma
--+
+-- Pustulosa_pustulosa__2 **
|
+-- Quadrula_quadrula
|
+-- Quadrula_verrucosa
Freeman et al., 2025 [Publ 4443], fig. 5 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 10 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Curole & Kocher, 2002 [Publ 2712], fig. 1a-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 3 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Curole & Kocher, 2002 [Publ 2712], fig. 1-F iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 3 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Curole & Kocher, 2005 [Publ 2682], fig. 1-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 11 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Chapman et al., 2008 [Publ 2667], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 20 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Curole & Kocher, 2002 [Publ 2712], fig. 1b-M iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 3 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Curole & Kocher, 2005 [Publ 2682], fig. 1 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 10 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Chase et al., 2018 [Publ 3847], fig. 4 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 38 | 1 | 1 | -- | +-- Cyclonaias
--+
+-- Pustulosa_pustulosa **
Breton et al., 2011 [Publ 3376], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 37 | 2 | 2 | -- | +----- Quadrula
--+
| +-- Cyclonaias
+--+
+-- Pustulosa_pustulosa **
Chapman et al., 2008 [Publ 2744], fig. 2 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 20 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_pustulosa **
Pieri et al., 2018 [Publ 3867], fig. 3 iT| tT | bi | riS | oT | roT | roS | iD | | 1 | 1 | 0 | 1 | 19 | 1 | 1 | -- | +-- Quadrula
--+
+-- Pustulosa_petrina **
|
