Palæos: Paleozoic		Permian Period
PALEOZOIC ERA		Permian: 2

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The Permian Period: 2

Phanerozoic Eon     Paleozoic Era         Cambrian Period         Ordovician Period         Silurian Period         Devonian Period         Carboniferous Period         Permian Period          Cisuralian Epoch (Early Permian)            Guadalupian Epoch (Middle Permian)            Lopingian Epoch (Late Permian)      Mesozoic Era           Triassic Period         Jurassic Period        Cretaceous Period     Cenozoic Era

Introduction Geography Stratigraphy Climate Sites Life Plants Vertebrates     Tetrapod Fossil Sites -- East Europe        Ufimian, Kazanian, and Tartarian        Stratigraphy of Tetrapods from East Europe

Vertebrates

The great diversity of near-shore and fresh water Chondrichthyans which characterized the Carboniferous began to decline during the Permian.  In the oceans, xenacanth sharks dominated until the Guadalupian, when they were replaced by hybodonts.   A few acanthodians also lingered into the Cisuralian.  Lungfishes and coelacanths were more diverse than they are today, but all of the other Sarcopterygian fishes had already become extinct. The Permian oceans were dominated by a diverse group of spiny-finned (actinopterygian) fishes, most of which had thick, heavy scales and rather basic jaw structures (if you look carefully, you can see this in the image of Palaeoniscus).  Neopterygians with more derived jaw structures probably began to appear by the end of the Lopingian.  

The increasing aridity of the Permian not only affected plants.  The tetrapods suffered as their swamps and pools shrunk and dried out.  Those surviving forms included big-headed temnospondyls two to three meters in length, as well as long-snouted forms (the archegosaurs) superficially resembling small crocodiles.  Many of the non-amniote reptilomorphs, such as anthracosaurs and embolomeres, continued in to the Permian. 

But it was the amniotes that took over as the dominant land animals, adapted to life on land (thanks to water-retentive dry skin and the amniotic egg).  Although there were a number of different types of amniotes, the largest and most diverse belonged to the Synapsida, which were ancestral to the mammals.

There were several distinct evolutionary dynasties of synapsids as the Permian progressed.

The first, the pelycosaur dynasty, included the large finbacks of the early Permian such as Dimetrodon,  Edaphosaurus, Ctenospondylus, and Secodontosaurus, all of which attained a lengths some 3 meters, as well a similar types that lacked a "sail".  The large dorsal "sails" were most certainly thermoregulatory devices that would heat up the animal in the cold morning, making it more active and giving it an advantage over it's more sluggish sail-less relatives.  These animals were limited to the equatorial tropics.

Following this was the Dinocephalian dynasty of the middle Permian (Guadalupian epoch).  The Dinocephalians were among the most primitive of the therapsids or "mammal-like reptiles".  Some grew to huge size (5 to 6 meters) with meter long heads full of wicked teeth (the name Dinocephalian means "fearsome head").  These creatures succeeded the Pelycosaurs, being both larger in size and more metabolically active.  There were several different types, the primitive anteosaurs (see Anteosaurus, left) being carnivores, and the ox-sized tapinocephalia being herbivores.  

The Dinocephalians all died out suddenly, perhaps as a result of unusual climatic factors, at the end of the Guadalupian.  The Therapsids that followed them were smaller, and more mammal-like.  Some may even evolved fur and the ability to control their temperatures metabolically.  These included the large gorgonopsians (the Permian equivalent of the "saber-toothed tiger"), the small to medium-sized Therocephalia, and the herbivorous dicynodonts.  These creatures had previously lived alongside the giant Dinocephalians, but came into their own when the latter had died out.  

In addition there were many types of non-syanapsid amniotes. The pareiasaurs, a group of big, armored herbivores probably related to turtles, reached enormous sizes (length up to 3 meters).   Smaller, lizard-like reptiles were probably common, but they are very poorly known.  These included the bolosaurs and procolophonids -- both also turtle relatives.  During the Permian, the crocodile-bird lineage (Archosauromorpha) had not yet diverged from the lizard-snake (Lepidosauromorpha) clade.  The primitive reptiles were represented by lizard-like captorhinids and basal diapsids.  The latter included a number of marine or amphibious forms: younginiforms, Claudiosaurus, and perhaps the earliest members of the ichthyosaur group.  

Image credits: Palaeoniscus from the Essex Rock & Mineral Society.  Deltavjatia from the University Museum of Zoology Cambridge.  

© MAK 2002.  Revised ATW050604.

Tetrapod Fossil Sites - East Europe

Ufimian, Kazanian, and Tartarian

The Russian late Permian strata are traditionally divided into the Ufimian (the earliest, immediately succeeding the Kungurian), the Kazanian, and the Tartarian (the latest, corresponding to the uppermost Permian deposits). Because of the importance of East European deposits in Permian stratigraphy (indeed, the Permian period is named after the Perm region in eastern Russia), these became the basis for a global standard. Unfortunately, the exact details of where each begins and ends is unfortunately somewhat obscure.

It turns out for example that the Tartarian ends some considerable time before the end of the Permian. According to Kozur 1998 the uppermost Tartarian is only earliest Wuchiapingian, which seems implausible in view of the advanced tetrapod fauna. Lozovsky 1998 in contrast indicates (at least from the diagrams) the Tartarian seems to stop at the middle Changhsingian, or thereabouts.

The fate of the Ufimian, which would generally be considered equivalent to perhaps the lower part of the Roadian, but is also sometimes considered belonging to a younger horizon, the Wordian (e.g. Gilmour and Morozova 1997), is even more unfortunate. On the basis of conodont zonation, Chernykh 2002 says:

"The upper Kazanian substage is most probably also Roadian, because Merrillina galeata, which is characteristic of higher stratigraphic horizons (Wordian) is absent in the Upper Kazanian of the Mid-Volga Region. Because the lower Kazanian correlates with the lower Roadian and I suggest the upper Kungurian correlates with the Cathedralian, it appears the Ufimian may lose its status as a stage."

I have therefore decided in these stratigraphic tables to exclude the Ufimian, equate the Kazanian with the Roadian, and the Tartarian with the Wordian through to Wuchiapingian

Stratigraphy of Tetrapods from East Europe

The following table, based mostly on information in Olson, 1962, King, 1990, Lozovsky, 1998, and Kurkin, 2001, catalogues the occurrence of tetrapods in East Europe. The five right-hand columns give the ranges for tetrapods found in those localities (see map, Permian localities in red)