BIOLOGY 622 – FALL 2014 BASAL AMNIOTA - STRUCTURE AND PHYLOGENY WEEK – 10 NYCTEROLETERIDAE AND PAREIASAUROMORPHA S. S. SUMIDA

Pareiasaurs include some of the largest and most dramatic of Late Paleozoic tetrapods. Though some are small, others got as large as cows. Recently, a somewhat obscure group of Russian forms known as the nycterolyterids have been suggested as close relatives of pareiasaurs, and Tsuji and Muller have brought them together in the larger taxon known as Pareiasauromorpha: Reptilia Eureptilia Parareptilia Ankyromorpha Pareiasauromorpha “Nycteroleteridae” Pareiasauridae

NYCTEROLETER PARAREPTILES The nycteroleters have been, until relatively recently, a somewhat obscure group of primarily Russian fORms form the middle to upper Permian. They have been obscure, in part, due to their documentation primarily in Cyrillic by Russian paleontologists. This has been ameliorated recently by the careful [re]descriptions of Macroleter and Emeroleter by Tsuji and colleagues.

If we look at recent global analyses of Parareptilia, the nycteroleters have been sometimes been considered a monophyletic group, and sometimes not. When considered monophyletic, the in- group relationships still have not been clear, and about the best we can say is that nycteroleters are more closely related tone another than anything else, but the intra-relationships remain largely unresolved. Note that in the following phylogeny from Tsuji et al (2012): The nycteroleters are considered a monophyletic group But, with the exception of Macroleter, Tsuji et al could only interpret the remaining five genera as an unresolved polychotomy.

Tsuji and colleagues defined the group with the following features: distinct emargination at the posterolateral edge of the skull concave, smooth depression in the temporal region extending across most of the squamosal and large parts of the quadratojugal remaining area of the temporal region is characterized by distinctive dermal sculpturing a well developed, laterally protruding rim at its dorsal margin of the otic notch formed by the overhanging supratemporal and postorbital. All of these features have been equated with the presence of a tympanic membrane in the notch to facilitate hearing in this group.

REPTILIA PARAREPTILIA PAREIASAUROMORPHA NYCEROLETERIDAE:

SURVEY OF NYCTEROLETER PARAREPTILES Macroleter and Emeroleter are the best known of the group. Notably, Emeroleter is the youngest of the group.

Macroleter Middle Permian Mezen River Basin, Arkhangel’sk Province, Russia and Oklahoma

Macroleter has been suggested by Reisz and Laurin (2001) to also be in Oklahoma based on the specimen below:

Emeroleter Late Permian Town of Kotel’nich, Kirov Province, Russia. Due to the work of Tsuji, Emeroleter is one of the best prepared and best known of the Russian nycteroleters.

DISTRIBUTION OF NYCTEROLETER PARAREPTILES All of the nycteroleters are from the middle or late Permian of Russia: In a present day map:

And in a map of the Late Permian, with Pangea still united: