Mikko's Phylogeny Archive

†Varanopidae

After Reisz, Dilkes & Berman, 1998, Reisz & Dilkes, 2003, Anderson & Reisz, 2004, Berman, Henrici, Sumida, Martens & Pelletier, 2014, and Brocklehurst, Reisz, Fernandez & Fröbisch, 2016

<==o †Varanopidae Romer & Price, 1940 sensu Reisz & Dilkes, 2003 [Varanopseidae (Romer & Price, 1940) Langston & Reisz, 1981; Varanopsidae]
   |?- †Basicranodon fortsillensis Vaughn, 1958
   |?- †Thrausmosaurus serratidens Fox, 1962
   |-- †Archaeovenator hamiltonensis Reisz & Dilkes, 2003
   `--+--+-- †Apsisaurus witteri Laurin, 1991
      |  `-- †Ascendonanus nestleri Spindler, Werneburg, Schneider, Luthardt, Annacker & Rößler, 2018; E.Perm. CEu.
      `--+-- †Pyozia mesenensis Anderson & Reisz, 2004
         `--+--o †Mesenosaurinae Romer, 1956 [Mycterosaurinae Reisz & Berman, 2001]
            |  |?-+-- †Anningia megalops Broom, 1927
            |  |  `-- †Microvaranops parentis Spindler, Werneburg, Schneider, Luthardt, Annacker & Rößler, 2018 [SAM-PK-K8305 (“Heleosaurus”)]; E.Perm. CEu.
            |  |-- †Heleosaurus scholtzi Broom, 1907
            |  `--+-- †Elliotsmithia longiceps Broom, 1937
            |     `--+-- †Mycterosaurus longiceps [Eumatthevia]
            |        `-- †Mesenosaurus romeri Efremov, 1938 [Mesenosauridae] [has been treated as a Diapsida incertae sedis]
            `--o †Varanopinae Romer, 1936 [Varanodontinae Reisz & Berman, 2001]
               |--+-- †Ruthiromia elcobriensis Eberth & Brinkman, 1983
               |  `--o †Aerosaurus Romer, 1937
               |     |-- †A. greenleeorum Romer, 1937
               |     `-- †A. wellesi Langston & Reisz, 1981
               `--+--+-- †Varanodon agilis Olson, 1965
                  |  `-- †Watongia meieri Olson, 1974
                  `--+-- †Tambacarnifex unguifalcatus Berman, Henrici, Sumida, Martens & Pelletier, 2014
                     `-- †Varanops brevirostris Williston, 1911 [Varanoops]

<==o †Varanopidae
   |--+-- †Apsisaurus witteri
   |  `-- †Ascendonanus nestleri
   `--+-- †Archaeovenator hamiltonensis
      `--+-- †Pyozia mesenensis
         `--o †Neovaranopsia
            |--o †Mesenosaurinae
            |  |--+-- †Mesenosaurus romeri
            |  |  `-- Varanopidae indet. [Cabarz, Thuringia]
            |  `--o †Afrothyra
            |     |-- †Elliotsmithia longiceps
            |     |-- †Heleosaurus scholtzi
            |     |-- BP/I/5678 [Alarmkraal farm]
            |     |-- SAM-PK-K10407 [La-de-da farm specimen]
            |     `--+-- †Anningia megalops
            |        `-- †Microvaranops parentis
            `--o †Varanodontinae
               |-- †Mycterosaurus longiceps
               `--+-- †Tambacarnifex unguifalcatus
                  `--+--+-- †“Aerosaurus” wellesi
                     |  `--+-- †Ruthiromia elcobriensis
                     |     `-- †Aerosaurus greenleeorum
                     `--+-- FMNH 644 [Richards Spur, cf. Varanops]
                        |-- †Varanops brevirostris
                        `--+-- †Varanodon agilis
                           `-- †Watongia meieri

Reference(s):

• Anderson, J. S. & Reisz, R. R., 2004: Pyozia mesenensis, a new small varanopid (Synapsida, Eupelycosauria) from Russia: "pelycosaur" diversity in the Middle Permian.
  –Journal of Vertebrate Paleontology: Vol. 24, #1, pp. 173-179
• Berman, D. S., Henrici, A. C., Sumida, S. S. & Martens, T., 2000: Redescription of Seymouria sanjuanensis (Seymouriamorpha) from the Lower Permian of Germany based on complete, mature specimens with a discussion of paleoecology of the Bromacker locality assemblage.
  –Journal of Vertebrate Paleontology: Vol. 20, #2, pp. 253-268
• Berman, D., Henrici, A., Sumida, S., Martens, T. & Pelletier, V., 2014: First European Record of a Varanodontine (Synapsida: Varanopidae): Member of a Unique Early Permian Upland Paleoecosystem, Tambach Basin, Central Germany.
  in Kammerer, C. F., Angielczyk, K. D. & Fröbisch, J., (eds.) 2014: Early Evolutionary History of the Synapsida.
  –Springer Netherlands, 2014, 69-86
• Botha-Brink, J. & Modesto, S. P., 2009: Anatomy and Relationships of the Middle Permian Varanopid Heleosaurus scholtzi Based on a Social Aggregation from the Karoo Basin of South Africa.
  –Journal of Vertebrate Paleontology: Vol. 29, #2, pp. 389-400 [doi: 10.1671/039.029.0209]
• Brocklehurst, N., Reisz, R. R., Fernandez, V. & Fröbisch, J., 2016: A Re-Description of 'Mycterosaurus' smithae, an Early Permian Eothyridid, and Its Impact on the Phylogeny of Pelycosaurian-Grade Synapsids.
  –Plos One: Vol. 11, #6, art.nr. e0156810 (1-27) [doi: 10.1371/journal.pone.0156810]
• Carroll, R. L., 1988: Vertebrate paleontology and evolution.
  –W. H. Freeman and company, New York, 1988, 698.
• Carroll, R. L., 1988: Appendix. 594-648.
  in Carroll, R. L. 1988: Vertebrate paleontology and evolution.
  –W. H. Freeman and company, New York, 1988, 698
• Evans, D. C., Maddin, H. C. & Reisz, R. R., 2009: A re-evaluation of sphenacodontid synapsid material from the lower Perrmian fissure fills near Richards Spur, Oklahoma.
  –Palaeontology: Vol. 52, #1, pp. 219-227 [doi: 10.1111/j.1475-4983.2008.00837.x]
• Kammerer, C. F. & Angielczyk, K. D., 2009: A proposed higher taxonomy of anomodont therapsids.
  –Zootaxa: #2018, pp. 1-24
• Langston, W. & Reisz, R. R., 1981: Aerosaurus wellesi, new species, a varanopseid mammal-like reptile (Synapsida: Pelycosauria) from the Lower Permian of New Mexico.
  –Journal of Vertebrate Paleontology: Vol. 1, #1, pp. 73-96
• Laurin, M. & Reisz, R. R., 1997-1998: Synapsida – Mammals and their extinct relatives.
  –iNet: The Tree of Life: http://tolweb.org/tree/eukaryotes/animals/chordata/synapsida.html
• Laurin , M. & Reisz, R. R., 1997: Autapomorphies of the main clades of synapsids.
  –iNet: The Tree of Life: http://tolweb.org/tree/eukaryotes/animals/chordata/synapsidlichen/synapsidasynapomorphies.html
• Reisz, R. R. & Dilkes, D. W., 2003: Archaeovenator hamiltonensis, a new varanopid (Synapsida: Eupelycosauria) from the Upper Carboniferous of Kansas.
  –Canadian Journal of Earth Sciencies: Vol. 40, pp. 667-678
• Reisz, R. R., Dilkes, D. W. & Berman, D. S., 1998: Anatomy and relationships of Elliotsmithia longiceps Broom, a small synapsid (Eupelycosauria: Varanopseidae) from the Late Permian of South Africa.
  –Journal of Vertebrate Paleontology: Vol. 18, #3, pp. 602-611
• Reisz, R. R., Laurin, M. & Marjanovic, D., 2010: Apsisaurus witteri from the Lower Permian of Texas: Yet Another Small Varanopid Synapsid, Not a Diapsid.
  –Journal of Vertebrate Paleontology: Vol. 30, #5, pp. 1628-1631 [doi: 10.1080/02724634.2010.501441]
• Reisz, R. R. & Modesto, S. P., 2007: Heleosaurus scholtzi from the Permian of South Africa: a varanopid synapsid, not a diapsid reptile.
  –Journal of Vertebrate Paleontology: Vol. 27, #3, pp. 734-739 [doi: 10.1671/0272-4634(2007)27[734:hsftpo]2.0.co;2]
• Spindler, F., Werneburg, R., Schneider, J. W., Luthardt, L., Annacker, V. & Rößler, R., 2018: First arboreal ‘pelycosaurs’ (Synapsida: Varanopidae) from the early Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny.
  –PalZ: in press [doi: 10.1007/s12542-018-0405-9]

Special thanks to:

Jos H. M. Dols, for gathering additional information and species of these animals.