Reconstruction of Thucydia mahoningensis, from Hernandez-Castillo et al. (2003).

Belongs within: Coniferophyta.

The Voltziales are early conifers known from the Late Carboniferous and Early Permian. Members were relatively small trees bearing plagiotropic lateral branches of determinate growth producing terminal cones (Rothwell et al. 2005).

<==Voltziales [Walchiaceae]
    |--+--Genoites patagonicaRMH-C05
    |  `--FerugliocladaceaeC93
    |       |--Ugartecladus genoensis Archangelsky & Cúneo 1987C93
    |       `--FerugliocladusRMH-C05
    |            |--F. patagonicus (Feruglio) Archangelsky & Cúneo 1987C93
    |            `--F. riojanum Archangelsky & Cúneo 1987C93
    `--+--Concholepis harrisiiRMH-C05
       |--Timanostrobus muravieviiRMH-C05
       `--+--Kungurodendron sharoviiRMH-C05
          `--+--Thucydia Hernandez-Castillo, Rothwell & Mapes 2001RMH-C05, H-CRM01 [Thucydiaceae]
             |    `--*T. mahoningensis Hernandez-Castillo, Rothwell & Mapes 2001H-CRM01
             `--+--Ernestiodendron filiciformeRMH-C05
                   |  |    |--O. junkeriH-CRM01
                   |  |    |--O. leonardii Florin 1964C93
                   |  |    `--O. visscheriH-CRM01
                   |  `--Voltziaceae [Cycadocarpidiacae, Swedenborgiaceae]C93
                   |       |  i. s.: Protodammara speciosa Hollick & Jeffrey 1909C93
                   |       |         Dectylolepis cyrptomerioides Hollick & Jeffrey 1909C93
                   |       |--VoltziaRMH-C05
                   |       |    |--V. heterophylla Brongniart 1835C93
                   |       |    |--V. hexagonaRMH-C05
                   |       |    `--V. hungaricaG31
                   |       `--+--AethophyllumRMH-C05
                   |          |    |--A. speciosum Schimper 1869C93
                   |          |    `--A. stipulareRMH-C05
                   |          `--MajonicaceaeC93
                   |               |--Dolmitia cittertiae Clement-Westerhof 1987C93
                   |               |--Majonica palina Clement-Westerhof 1987RMH-C05, C93
                   |               `--PseudovoltziaC93
                   |                    |--P. liebeana (Geinitz) Florin 1927C93 [=Voltzia liebeanaH-CRM01]
                   |                    `--P. sjerpii Clement-Westerhof 1987C93
                   `--Emporiaceae [Bartheliaceae]RMH-C05
                        |  i. s.: PotonieisporitesRMH-C05
                        |           |--*P. novicusRM01
                        |           |--P. granulatusRM01
                        |           `--P. neglectusRM01
                        |--Hanskerpia Rothwell, Mapes & Hernandez-Castillo 2005RMH-C05
                        |    `--*H. hamiltonensis Rothwell, Mapes & Hernandez-Castillo 2005RMH-C05
                        `--+--Barthelia Rothwell & Mapes 2001RMH-C05, RM01
                           |    `--*B. furcata Rothwell & Mapes 2001RM01
                           `--+--Emporia Mapes & Rothwell 2003RMH-C05, MR03
                              |    `--*E. lockardii (Mapes & Rothwell) Mapes & Rothwell 2003 (see below for synonymy)MR03
                              `--Utrechtiaceae [Lebachiaceae]RMH-C05
                                   |--Walchianthus cylindraceusRM01
                                   |--Otovicia hypnoides [=Lebachia hypnoides]RMH-C05
                                   `--Utrechtia Mapes & Rothwell in Rothwell & Mapes 2003RMH-C05, RM03 (see below for synonymy)
                                        |--*U. floriniformis Mapes & Rothwell in Rothwell & Mapes 2003RM03 (see below for synonymy)
                                        |--‘Lebachia’ americanaRM01
                                        |--‘Walchia’ arnhardtii [=Ernestodendron arnhardtii]H-CRM01
                                        |--‘Walchia’ crassusRM01
                                        |--‘Walchia’ filiformis G31
                                        |--‘Lebachia’ frondosaH-CRM01
                                        |--‘Lebachia’ garnettensisH-CRM01
                                        |--‘Walchia’ germanica [=Ernestodendron germanica]H-CRM01
                                        |--‘Lebachia’ goeppertianaH-CRM01
                                        |--‘Lebachia’ laxifoliaH-CRM01
                                        |--‘Lebachia’ parvifoliaH-CRM01
                                        |--‘Walchia’ schlotheimii [=Lebachia schlotheimii]H-CRM01
                                        `--‘Lebachia’ speciosaH-CRM01

Voltziales incertae sedis:
    |--L. gouldiiH-CRM01
    `--L. grandeuryiH-CRM01
  Ullmannia Göppert 1850H-CRM01, C93 [Ullmanniaceae]
    |--U. biarmicaG31
    |--U. bronnii Göppert 1850C93
    |--U. caulerpaK64
    |--U. frumentaria Göppert 1850C93
    |--U. geinitziG31
    `--U. selaginoidesG31
  Walchiostrobus [incl. Thuringiostrobus]H-CRM01
    |--W. gothanii [=Lebachia gothanii]H-CRM01
    `--W. lodevensisH-CRM01
  Batenburgia sakmaricaH-CRM01
  Maculostrobus willsi Townrow 1962W70

*Emporia lockardii (Mapes & Rothwell) Mapes & Rothwell 2003 [=Lebachia lockardii Mapes & Rothwell 1984]MR03

Utrechtia Mapes & Rothwell in Rothwell & Mapes 2003RMH-C05, RM03 [incl. HermitiaRM03, Lebachia (nom. illeg.)RM03]

Utrechtia floriniformis Mapes & Rothwell in Rothwell & Mapes 2003RM03 [incl. *Lebachia piniformisH-CRM01, *Walchia piniformis (n. d.)H-CRM01]

*Type species of generic name indicated


[C93] Cleal, C. J. 1993. Gymnospermophyta. In: Benton, M. J. (ed.) The Fossil Record 2 pp. 795–808. Chapman & Hall: London.

[G31] Grabau, A. W. 1931. The Permian of Mongolia: A report on the Permian fauna of the Jisu Honguer limestone of Mongolia and its relations to the Permian of other parts of the world. American Museum of Natural History: New York.

[H-CRM01] Hernandez-Castillo, G. R., G. W. Rothwell & G. Mapes. 2001. Thucydiaceae fam. nov., with a review and reevaluation of Paleozoic walchian conifers. International Journal of Plant Science 162 (5): 1155–1185.

[K64] Kirkby, J. W. 1864. On some fossil fishes from the Permian limestone of Fulwell, near Sunderland. Report of the British Association for the Advancement of Science 33 (Notices and Abstracts): 82.

[MR03] Mapes, G., & G. W. Rothwell. 2003. Validation of the names Emporiaceae, Emporia, and Emporia lockardii. Taxon 52: 327–328.

[RM01] Rothwell, G. W., & G. Mapes. 2001. Barthelia furcata gen. et sp. nov., with a review of Paleozoic coniferophytes and a discussion of coniferophyte systematics. International Journal of Plant Sciences 162 (3): 637–667.

[RM03] Rothwell, G. W., & G. Mapes. 2003. Validation of the names Utrechtiaceae, Utrechtia, and Utrechtia floriniformis. Taxon 52: 329–330.

[RMH-C05] Rothwell, G. W., G. Mapes & G. R. Hernandez-Castillo. 2005. Hanskerpia gen. nov. and phylogenetic relationships among the most ancient conifers (Voltziales). Taxon 54 (3): 733–750.

[W70] Wills, L. J. 1970. The Triassic succession in the central Midlands in its regional setting. Quarterly Journal of the Geological Society of London 126: 225–283.

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