species Pustulosa pustulosa (Lea, 1831): synonymy

Obliquaria (Quadrula) bullata Rafinesque, 1820

(homonym of Obliquaria (Quadrula) flexuosa var. bullata Rafinesque, 1820)
Rafinesque, 1820. Ann. Gén. Sci. Phys. 5 (13): 307 [also as ‘U. bullata’].
Rafinesque, 1864. [in] Binney & Tryon, reprint: 51.
NEOTYPE ANSP_20250, Kentucky River.

Quadrula pustulosa pernodosa — Vanatta (1915)
Quadrula pustulosa — Ortmann (1918), Ortmann & Walker (1922), Burch (1975), R.I. Johnson (1980), Parmalee & Bogan (1998), Williams et al. (2008)
Quadrula (Bullata) pustulosa — Frierson (1927)
Pustulosa bullata — Morrison (1966)
Quadrula (Pustulosa) pustulosa pustulosa — Haas (1969)
Quadrula bullata — Morrison (1969)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Obliquaria (Quadrula) retusa Rafinesque, 1820

(renamed Unio premorsus Rafinesque, 1831)
Rafinesque, 1818. Amer. Mon. Mag. 3: 355 [as ‘Potamilus retusus’]. [nomen nudum]
Rafinesque, 1819. Jour. Phys. Chim. Hist. Nat. 88: 427 [as ‘U. (Obovaria) retusa’]. [nomen nudum]
Rafinesque, 1820. Ann. Gén. Sci. Phys. 5 (13): 306, pl. 81, figs. 19-20 [also as ‘Unio retusa’]. [nomen oblitum]
Rafinesque, 1864. [in] Binney & Tryon, reprint: 22, 29, 50.
NEOTYPE ANSP_20220, Green River.

Quadrula pustulosa — Vanatta (1915), Ortmann & Walker (1922), Williams et al. (2008)
Obliquaria retusa — Burch (1975)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio verrucosa Valenciennes, 1827

(homonym of Unio verrucosus Barnes, 1823)
Valenciennes, 1827. Receuil Observ. Zool. 2(11-12): 231, pl. 53, fig. 2.

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa — Simpson (1914)
Quadrula (Bullata) pustulosa — Frierson (1927)
Quadrula (Pustulosa) pustulosa pustulosa — Haas (1969)
Quadrula pustulosa — Parmalee & Bogan (1998)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio premorsus Rafinesque, 1831

(new name for Obliquaria (Quadrula) retusa Rafinesque, 1820)
Rafinesque, 1831. Continuation: 4.
Rafinesque, 1864. [in] Binney & Tryon, reprint: 77.

Pleurobema premorsa — Morrison (1969)
Quadrula pustulosa — Williams et al. (2008)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio pustulosus Lea, 1831

Lea, 1831. Trans. Amer. Phil. Soc. 4 [O. 1]: 76 [86], pl. 7, fig. 7.
SYNTYPES ANSP_43058, Ohio.

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Unio (s.s.) pustulosus — Call (1900)
Quadrula pustulosa — Ortmann (1909), Ortmann (1909), Ortmann (1909), Ortmann (1910), Ortmann (1911), ... Jones (2015), Inoue et al. (2018), Marshall et al. (2018), Klymus et al. (2020), Gasparini et al. (2020)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa — Simpson (1914)
Unio pustulosus — de Gregorio (1914)
Quadrula (Bullata) pustulosa — Frierson (1927)
Quadrula (s.s.) pustulosus — F.C. Baker (1928)
Quadrula (Pustulosa) pustulosa pustulosa — Haas (1969)
Quadrula bullata — Morrison (1969)
Quadrula pustulosa pustulosa — Turgeon et al. (1988), Williams et al. (1993), Turgeon et al. (1998), Lydeard et al. (1999), Watters et al. (2009), ITIS (2012)
Amphinaias pustulosa — Graf & Cummings (2007), Cummings & Graf (2010), IUCN (2011), Roe & Boyer (2015)
Rotundaria pustulosa — Campbell & Lydeard (2012)
Cyclonaias pustulosa — Williams et al. (2017), Johnson et al. (2018), Lopes-Lima et al. (2019), Graf & Cummings (2021), Bieler (2021), FMCS (2021), de Moulpied et al. (2022), Hopper et al. (2023), Freeman et al. (2025)
Pustulosa pustulosa — Neemuchwala et al. (2023), FMCS (2025), Campbell & Lydeard (2026)
Pustulosa (Cyclonaias) pustulosa — Rodriguez et al. (2023)

Unio nodulosus Say, 1834

(misidentified reference to Mya nodulosa Wood, 1815)
Say, 1834. Amer. Conch. 6: ??.

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa — Simpson (1914)
Quadrula pustulosa — Utterback (1915), Oesch (1984), Howells et al. (1996), Parmalee & Bogan (1998), Williams et al. (2008)
Quadrula (Bullata) pustulosa — Frierson (1927)
Quadrula (Pustulosa) pustulosa pustulosa — Haas (1969)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio prasinus Conrad, 1834

Conrad, 1834. New Fresh-Water Shells: 44, pl. 3, fig.1.
Conrad, 1837. Monogr. Fam. Unionidae 1(9): 79, pl. 44, fig. 2.
Conrad, 1853. Proc. Acad. Nat. Sci. 6: 255.

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa — Simpson (1914)
Quadrula pustulosa prasina — Ortmann & Walker (1922)
Quadrula (Bullata) pustulosa-prasina — Frierson (1927)
Quadrula (s.s.) pustulosus prasina — F.C. Baker (1928)
Quadrula (Pustulosa) pustulosa prasina — Haas (1969)
Quadrula pustulosa — Burch (1975), Parmalee & Bogan (1998), Williams et al. (2008)
Quadrula quadrula — R.I. Johnson (1980)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio schoolcraftensis Lea, 1834

Lea, 1834. Trans. Amer. Phil. Soc. 5 [O. 1]: 37 [149], pl. 3, fig. 9.

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa var. schoolcraftensis — Simpson (1914)
Quadrula pustulosa schoolcraftensis — Utterback (1915)
Quadrula pustulosa prasina — Ortmann & Walker (1922)
Quadrula (Bullata) pustulosa-prasina — Frierson (1927)
Quadrula (s.s.) pustulosus prasina — F.C. Baker (1928)
Quadrula (Pustulosa) pustulosa prasina — Haas (1969)
Quadrula pustulosa — R.I. Johnson (1980), Parmalee & Bogan (1998), Williams et al. (2008)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019), Bieler (2021)

Unio mortoni Conrad, 1835

Conrad, 1835. Monogr. Fam. Unionidae 1(1): 11, pl. 6, fig. 1.
Conrad, 1853. Proc. Acad. Nat. Sci. 6: 252.
HOLOTYPE ANSP_10287A, Bayou Teche, Louisiana.

Quadrula (sec. Theliderma) mortoni — Simpson (1900)
Quadrula mortoni — Ortmann (1912), Vidrine (1993), Howells et al. (1996), Serb et al. (2003), Bogan (2010), Pfeiffer et al. (2016), Marshall et al. (2018)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) mortoni — Simpson (1914)
Quadrula (Bullata) pustulosa-mortoni — Frierson (1927)
Quadrula (Pustulosa) pustulosa mortoni — Haas (1969)
Quadrula pustulosa mortoni — Turgeon et al. (1988), Williams et al. (1993), Turgeon et al. (1998), ITIS (2012), Szumowski et al. (2012)
Quadrula pustulosa — Parmalee & Bogan (1998), Williams et al. (2008), Haag (2012)
Cyclonaias mortoni — Williams et al. (2017)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019), FMCS (2021)
Pustulosa (Cyclonaias) pustulosa — Rodriguez et al. (2023)
Pustulosa pustulosa — FMCS (2025)

Unio bullatus ‘Rafinesque’ of Authors

(reference to Obliquaria (Quadrula) bullata Rafinesque, 1820)
‘Rafinesque’ of Authors.
Conrad, 1838. Monogr. Fam. Unionidae 1(10): 82, pl. 45, fig. 2.
Lapham, 1852. Trans. WI Agri. Soc. 2: 369.
Reeve, 1864. Conch. Icon. 16: pl. 12, sp. 43.

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa — Simpson (1914)
Quadrula (s.s.) pustulosus — F.C. Baker (1928)
Quadrula pustulosa pustulosa — Watters et al. (2009)

Unio dorfeuillianus Lea, 1838

(correct name for Unio dorfuillianus Lea, 1836)
Lea, 1836. Synopsis Fam. Naiades: addenda [as ‘dorfuillianus’]. [nomen nudum]
Lea, 1838. Trans. Amer. Phil. Soc. 6 [O. 2]: 73, pl. 17, fig. 54.
HOLOTYPE USNM_84215, Wabash River, Ind.

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Unio (s.s.) pustulosus — Call (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa — Simpson (1914)
Quadrula (Bullata) nodulata — Frierson (1927)
Quadrula (s.s.) pustulosus — F.C. Baker (1928)
Quadrula pustulosa — Murray & Leonard (1962), Oesch (1984), Howells et al. (1996), Parmalee & Bogan (1998), Williams et al. (2008)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio turgidus Lea, 1838

Lea, 1836. Synopsis Fam. Naiades: 16. [nomen nudum]
Lea, 1838. Trans. Amer. Phil. Soc. 6 [O. 2]: 11, pl. 5, fig. 11.
HOLOTYPE USNM_84277, near New Orleans, La.

Quadrula (sec. Theliderma) mortoni — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) mortoni — Simpson (1914)
Quadrula (Bullata) pustulosa-mortoni — Frierson (1927)
Quadrula (Pustulosa) pustulosa mortoni — Haas (1969)
Quadrula mortoni — Vidrine (1993), Howells et al. (1996)
Quadrula pustulosa — Parmalee & Bogan (1998), Williams et al. (2008)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio nodiferus Conrad, 1841

Conrad, 1841. Proc. Acad. Nat. Sci. 1: 19.
Conrad, 1850. Jour. Acad. Nat. Sci. 1: 277, pl. 38, figs. 4, 8.
Conrad, 1853. Proc. Acad. Nat. Sci. 6: 253.
LECTOTYPE ANSP_10285, Jackson, La.

Quadrula (sec. Theliderma) nodifera — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) nodifera — Simpson (1914)
Quadrula (Bullata) pustulosa-mortoni — Frierson (1927)
Quadrula (Pustulosa) pustulosa mortoni — Haas (1969)
Quadrula pustulosa — Burch (1975)
Quadrula mortoni — Vidrine (1993), Howells et al. (1996)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio pernodosus Lea, 1845

Lea, 1845. Proc. Amer. Phil. Soc. 4: 163.
Lea, 1848. Trans. Amer. Phil. Soc. 10 [O. 4]: 71 [45], pl. 3, fig. 8.
SYNTYPE USNM_84298, N.C.

Quadrula (sec. Theliderma) pustulosa var. pernodosa — Simpson (1900)
Quadrula pernodosa — Ihering (1901)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa var. pernodosa — Simpson (1914)
Quadrula pustulosa pernodosa — Vanatta (1915)
Quadrula pustulosa — Ortmann (1918), Ortmann & Walker (1922), Burch (1975), Parmalee & Bogan (1998), Williams et al. (2008)
Quadrula (Bullata) pustulosa-asperata — Frierson (1927)
Quadrula (Pustulosa) pustulosa pernodosa — Haas (1969)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio aureus Lea, 1859

Lea, 1859. Proc. Acad. Nat. Sci. 11: 112.
Lea, 1862. Jour. Acad. Nat. Sci. 5 [O. 9]: 195 [17], pl. 26, fig. 264.
HOLOTYPE USNM_84572, Texas.

Quadrula (sec. Theliderma) aurea — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) aurea — Simpson (1914)
Quadrula (Bullata) aureus — Frierson (1927)
Quadrula (Pustulosa) aurea — Haas (1969)
Quadrula aurea — Burch (1975), Turgeon et al. (1988), Williams et al. (1993), Howells et al. (1996), Turgeon et al. (1998), Serb et al. (2003), Bogan (2010), ITIS (2012), Szumowski et al. (2012), Haag (2012)
Amphinaias aurea — Graf & Cummings (2007), Cummings & Graf (2010)
Cyclonaias aurea — Williams et al. (2017)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019), Graf & Cummings (2021), FMCS (2021)
Pustulosa (Cyclonaias) pustulosa — Rodriguez et al. (2023)
Pustulosa pustulosa — FMCS (2025)

Unio houstonensis Lea, 1859

Lea, 1859. Proc. Acad. Nat. Sci. 11: 155.
Lea, 1860. Jour. Acad. Nat. Sci. 4 [O. 8]: 358 [40], pl. 60, fig. 183.
HOLOTYPE USNM_85768, Rutersville, Texas.

Quadrula (sec. Theliderma) houstonensis — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) houstonensis — Simpson (1914)
Quadrula (Bullata) houstonensis — Frierson (1927)
Quadrula houstonensis — Turgeon et al. (1988), Williams et al. (1993), Howells et al. (1996), Turgeon et al. (1998), Bogan (2010), Breton et al. (2011), ITIS (2012), Haag (2012), Pfeiffer et al. (2016), Chase et al. (2018)
Pleurobema riddelli — Vidrine (1993)
Amphinaias houstonensis — IUCN (1996), Graf & Cummings (2007), Cummings & Graf (2010)
Cyclonaias houstonensis — Williams et al. (2017)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019), Graf & Cummings (2021), FMCS (2021)
Pustulosa (Cyclonaias) pustulosa — Rodriguez et al. (2023)
Pustulosa pustulosa — FMCS (2025)

Unio pauperculus Lea, 1861

Lea, 1861. Proc. Acad. Nat. Sci. 13: 39.
Lea, 1862. Jour. Acad. Nat. Sci. 5 [O. 8]: 99 [103], pl. 15, fig. 247.
HOLOTYPE USNM_85763, Columbus, Miss.

Quadrula (sec. Theliderma) paupercula — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) paupercula — Simpson (1914)
Quadrula (Bullata) pustulosa-paupercula — Frierson (1927)
Quadrula (Pustulosa) pustulosa paupercula — Haas (1969)
Quadrula asperata — Williams et al. (2008)
Cyclonaias pustulosa — Lopes-Lima et al. (2019)

Unio refulgens Lea, 1868

Lea, 1868. Proc. Acad. Nat. Sci. 20: 145.
Lea, 1868. Jour. Acad. Nat. Sci. 6 [O. 12]: 317 [77], pl. 51, fig. 130.
FIGURED SYNTYPE USNM_84290, Enterprise, Lauderdale Co., Miss. [Alabama].

Quadrula (sec. Theliderma) refulgens — Simpson (1900)
Quadrula refulgens — Ortmann (1912), Turgeon et al. (1988), Williams et al. (1993), Vidrine (1993), Turgeon et al. (1998), ... Bogan (2010), ITIS (2012), Szumowski et al. (2012), Haag (2012), Inoue et al. (2018)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) refulgens — Simpson (1914)
Quadrula (Bullata) pustulosa-refulgens — Frierson (1927)
Quadrula (Pustulosa) pustulosa refulgens — Haas (1969)
Quadrula pustulosa — Burch (1975), R.I. Johnson (1980)
Amphinaias refulgens — IUCN (1996), Graf & Cummings (2007), Cummings & Graf (2010)
Rotundaria refulgens — Campbell & Lydeard (2012), Campbell & Lydeard (2012)
Cyclonaias refulgens — Williams et al. (2017)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019), Graf & Cummings (2021), FMCS (2021)
Cyclonaias (Quadrula) refulgens — Porto-Hannes et al. (2021)
Pustulosa (Cyclonaias) pustulosa — Rodriguez et al. (2023)
Pustulosa pustulosa — FMCS (2025)

Unio sphaericus Lea, 1868

Lea, 1868. Proc. Acad. Nat. Sci. 20: 145 [as ‘sphæricus’].
Lea, 1868. Jour. Acad. Nat. Sci. 6 [O. 12]: 319 [79], pl. 51, fig. 132 [as ‘sphæricus’].
FIGURED SYNTYPE USNM_84271, Pearl River, Jackson, Miss.

Quadrula (sec. Theliderma) sphærica — Simpson (1900)
Quadrula sphaerica — Ortmann (1912)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) sphærica — Simpson (1914)
Quadrula pustulosa — Ortmann (1918), Burch (1975)
Quadrula (Bullata) pustulosa-refulgens — Frierson (1927)
Quadrula mortoni — Vidrine (1993), Howells et al. (1996)
Cyclonaias pustulosa — Johnson et al. (2018), Lopes-Lima et al. (2019)

Unio vallatus Lea, 1868

Lea, 1868. Proc. Acad. Nat. Sci. 20: 145.
Lea, 1868. Jour. Acad. Nat. Sci. 6 [O. 12]: 315 [75], pl. 50, fig. 128.
HOLOTYPE (MONOTYPY) USNM_84275, Alabama R., Ala.

Quadrula (sec. Theliderma) vallata — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) vallata — Simpson (1914)
Quadrula (Bullata) pustulosa-vallata — Frierson (1927)
Quadrula (Pustulosa) pustulosa vallata — Haas (1969)
Quadrula pustulosa — Burch (1975)
Quadrula asperata — Williams et al. (2008)
Cyclonaias pustulosa — Lopes-Lima et al. (2019)

Unio schoolcraftii ‘Lea’ of Authors

(error for Unio schoolcraftensis Lea, 1834)
‘Lea’ of Authors.
Lea, 1870. Synopsis Fam. Naiades: 33.
B.H. Wright, 1888. Check List N. Amer. Unionidae etc.: [5].

Quadrula (sec. Theliderma) pustulosa — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) pustulosa var. schoolcraftensis — Simpson (1914)
Quadrula (Bullata) pustulosa-prasina — Frierson (1927)
Quadrula (Pustulosa) pustulosa prasina — Haas (1969)
Quadrula pustulosa — Vidrine (1993), Parmalee & Bogan (1998), Williams et al. (2008)
Quadrula pustulosa pustulosa — Watters et al. (2009)
Cyclonaias pustulosa — Bieler (2021)

Unio cahabensis Lea, 1871

Lea, 1871. Proc. Acad. Nat. Sci. 23: 190.
Lea, 1874. Jour. Acad. Nat. Sci. 8 [O. 13]: 17 [21], pl. 5, fig. 14.
FIGURED SYNTYPE USNM_84204, Cahaba R., Ala.

Quadrula (sec. Theliderma) cahabensis — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) cahabensis — Simpson (1914)
Quadrula (Bullata) pustulosa-vallata — Frierson (1927)
Quadrula (Pustulosa) pustulosa vallata — Haas (1969)
Quadrula pustulosa — Burch (1975)
Quadrula asperata — Williams et al. (2008)
Cyclonaias pustulosa — Lopes-Lima et al. (2019)

Quadrula pustulosa asperatus Utterback, 1915

(misidentified reference to Unio asperatus Lea, 1861)
‘Lea’ Utterback, 1915. Amer. Midl. Nat. 4(4): 134, pl. 17, figs. 42a-b.

Quadrula pustulosa — Parmalee & Bogan (1998)
Quadrula pustulosa pustulosa — Watters et al. (2009)

Unio petrinus Frierson, 1927

(intro'd as syn of Unio aureus Lea, 1859)
‘Call (Mss)’ Frierson, 1927. Check List N. Amer. Naiades: 49.

Quadrula (Bullata) aureus — Frierson (1927)
Cyclonaias pustulosa — Johnson et al. (2018)

Unio verrucosus albus Say of Authors

(misidentified reference to Unio verrucosus albus Hildreth, 1828)
Say of Authors.

Unio dorfuillianus Lea, 1836

(incorrect original spelling of Unio dorfeuillianus Lea, 1838)
Lea, 1836. Synopsis Fam. Naiades: addenda. [nomen nudum]

Unio postulosus Lapham, 1860

(error for Unio pustulosus Lea, 1831)
‘Lea’ Lapham, 1860. Proc. Acad. Nat. Sci. 12: 155.

Margaron (Unio) spaericus Lea, 1870

(error for Unio sphaericus Lea, 1868)
‘Lea’ Lea, 1870. Synopsis Fam. Naiades: 33 [as ‘spæricus’].

Unio nodiferous Singley, 1893

(error for Unio nodiferus Conrad, 1841)
‘Conrad’ Singley, 1893. Geol. Surv. Texas (1892): 319.

Unio turgidis Singley, 1893

(error for Unio turgidus Lea, 1838)
‘Lea’ Singley, 1893. Geol. Surv. Texas (1892): 319.

Unio pustulosus Call, 1895

(reference to Unio pustulosus Lea, 1831)
‘Lea’ Call, 1895. Trans. Acad. Sci. St. Louis 7: 42, pl. 15, figs. 3-4.

Quadrula (sec. Theliderma) refulgens — Simpson (1900)
Quadrula (sec. Theliderma) (Group of Q. pustulosa) refulgens — Simpson (1914)