Monday, September 1, 2014

Meet the Chroniosuchians

Chroniosaurus dongusensis measured about 30 cm in length.
Chroniosuchians form a very obscure group of small, superficially croc-like four-legged animals that lived during the Late Permian period and survived until the Middle and perhaps Late Triassic periods. Most species are from the Late Permian (Late Tatarian regional stage) of Russia, but they are also known from China, Germany and Kyrgyzstan. They were aquatic or semi-aquatic fish-eating predators characterized by a distinct row of interconnected “butterfly-shaped” bony plates or scutes (osteoderms) covering their back and part of their tail, one for each vertebra. They are traditionally subdivided into two families, the more primitive Chroniosuchidae from the Late Permian with one representative in the Triassic and the Bystrowianidae from the Late Permian to the Middle Triassic. The exact affinities of the Chroniosuchians are quite uncertain. They are usually placed among non-amniote  reptiliomorphs but more recent studies make them “stem tetrapods”, so not reptiles and not quite amphibians but something a bit more primitive. The eel-like embolomeres appear to have been their closest relatives.

Madygenerpeton pustulatus from Kyrgyzstan.
Among the Chroniosuchids, Chroniosaurus dongusensis Tverdochlebova, 1972 from the Late Permian of the Orenburg province of Russia is probably the best known species, with fossils from many individuals including one near complete articulated specimen. Chroniosuchus paradoxus Vjuschkov, 1957 also from the Orenburg province but a bit younger, differs in the taller shape of its skull. Madygenerpeton pustulatus Schoch et al., 2010 from the Middle or Late Triassic of Kyrgyzstan appears to be the basalmost member of the group despite its late occurrence. The type specimen is a nearly complete skull showing a broad snout quite different from the two previous species. Jarilinus mirabilis (Vjuschkov, 1957) from the Novgorod province and Uralerpeton  tverdochlebovae Golubev, 1998 from the Vladimir Province were quite large species for the group, the reconstructed skull of the former measuring 20 cm and of the latter some 55 cm. The others three species, Chroniosaurus levis Golubev, 1908, Chroniosuchus lichaveri (Riabinin, 1962) and Suchonica vladimiri Golubev, 1999, all from the Vologda Province of Russia, are only known from fragmentary remains.

Chroniosuchus paradoxus.
The other family, the Bystrowianids, are much less known, their fossil remains consisting mainly on isolated armor scutes. Late permian species include Bystrowiana permira Vjuschkov, 1957 from the Vladimir Province of Russia, Bystrowiana sinica Young, 1979, Dromotectum largum Liu et al., 2014 and Jinyuanitectum flatum Liu et al., 2014, the last three from the Henan Province of China. Early Triassic species are Axitectum vjushkovi Shishkin & Novikov, 1992 from the Novgorod province, Axitectum georgi Novikov & Shishkin, 2000 from the Kirov province, Dromotectum spinosum Novikov & Shishkin, 1996 from the Orenburg Province. Middle Triassic forms are Synesuchus muravjevi Novikov & Shishkin, 2000 from the Komi Republic and Bystrowiella schumanni Witzmann et al., 2008 from Germany.


Buchwitz, M., Foth, C., Kogan, I., & Voigt, S. (2012). On the use of osteoderm features in a phylogenetic approach on the internal relationships of the Chroniosuchia (Tetrapoda: Reptiliomorpha). Palaeontology, 55(3), 623–640.
Golubev, V. K. (1998). Narrow-armored Chroniosuchians from the Late Permian of Eastern Europe. Paleontological Journal, 32(3), 278–287.
Golubev, V. (1998). Revision of the Late Permian chroniosuchians (Amphibia, Anthracosauromorpha) from Eastern Europe. Paleontological Journal, 32(4), 390–401.
Golubev, V. K. (1999). A New Narrow-Armored Chroniosuchian from the Upper Permian of Eastern Europe. Paleontological Journal, 33(2), 166–173.
Klembara, J., Clack, J. a., & Čerňanský, A. (2010). The anatomy of palate of Chroniosaurus dongusensis (Chroniosuchia, Chroniosuchidae) from the Upper Permian of Russia. Palaeontology, 53(5), 1147–1153.
Liu, J., Xu, L., Jia, S.-H., Pu, H.-Y., & Liu, X.-L. (2014). The Jiyuan tetrapod fauna of the Upper Permian of China — 2 . stratigraphy , taxonomical review , and correlation. Vertebrata PalAsiatica, 52(3), 328–339.
Schoch, R., Voigt, S., & Buchwitz, M. (2010). A chroniosuchid from the Triassic of Kyrgyzstan and analysis of chroniosuchian relationships. Zoological Journal of the Linnean Society, 160, 515–530.
Witzmann, F., Schoch, R. R., & Maisch, M. W. (2008). A relict basal tetrapod from Germany: first evidence of a Triassic chroniosuchian outside Russia. Die Naturwissenschaften, 95(1), 67–72.