New papers on Kyrgyzsaurus, Madygenerpeton, and Longisquama

Alifanov, V. R. and E. N. Kurochkin. 2011..Kyrgyzsaurus bukhanchenkoi gen. et sp. nov., a new reptile from the Triassic of southwestern Kyrgyzstan. Paleontological Journal 45(6):639-647.
[DOI: 10.1134/S0031030111060025] [link]

Description of a reptile fossil with skin preservation discovered in 2006. Comes form the same locality as Sharovipteryx and Longisquama. The authors interpret the specimen as a member of drepanosaurs, a Late Triassic group of archosauromorphs. This paper represents one of the last contributions of the Russian palaeornithologist Evgenii N. Kurochkin who passed away recently.

Buchwitz, M., C. Foth, I. Kogan, and S. Voigt. 2012 in press. On the use of osteoderm features in a phylogenetic approach on the internal relationships of the Chroniosuchia (Tetrapoda: Reptiliomorpha). Palaeontology. [DOI: 10.1111/j.1475-4983.2012.01137.x] [link]

Includes a graphic reconstruction of Madygenerpeton (drawing by Frederik Spindler).

Buchwitz, M. and S. Voigt. 2012 in press. The dorsal appendages of the Triassic reptile Longisquama insignis: reconsideration of a controversial integument type. Paläontologische Zeitschrift.
[DOI: 10.1007/s12542-012-0135-3] [Link]

More thorough description/ graphic documentation compared to Voigt et al.(2009) and considers some aspects of diapsid skin evolution.

A palaeodictyopteran and other relics from Madygen

Béthoux, O., S. Voigt, and J. W. Schneider. 2010. A Triassic palaeodictyopteran from Kyrgyzstan. Palaeodiversity 3: 9-13. [pdf 1.5 Mb]

Despite the substantial collection and study of insect fossils from the Madygen Formation (see overview in Shcherbakov 2008a) there are still unkown elements of the entomofauna left. Béthoux et al. (2010) describe a wing of a not yet reported group of insects from lacustrine shales of the northwestern ouctrop area of the Madygen Fm. (which also yielded Sharovipteryx and Longisquama).

Ruling out all alternatives on the basis of wing venation data, they come to the conclusion that reliquia spec. nov. was a late member of Palaeodictyoptera, an order-rank group according to conventional classification schemes that was previously thought to have died out during the Middle or Late Permian.

Béthoux et al. suggest that the disappearance of ancient insect groups in equatorial realms is linked to the Late Paleozoic aridisation in these areas that triggered the migration to wetter higher latitude ecosystems, such as the Madygen lake environment. The relatively late occurence of paleodictyopterans in Madygen is also in agreement with Shcherbakov's (2008b) hypothesis that the renewal of Triassic entomofaunas was asynchronous, starting in the lower latitudes and spreading to the higher latitudes.

Other Madygen relics?

Apart from modern groups, such as dipterans and hymenopterans among insects as wells as lissamphibians and archosaurs among tetrapods there are further relict forms, such as the choniosuchian Madygenerpeton or the basal cynodont Madysaurus. As hinted by Béthoux et al. the question to what degree and why Madygen functioned as a refugium is still to be answered.

Chroniosuchia: Paper on osteoderm histology in online preview

...my first experience with bone histology:

Buchwitz, M., Witzmann, F., Voigt, S. & Golubev, V. in press. Osteoderm microstructure indicates the presence of a crocodylian-like trunk bracing system in a group of armoured basal tetrapods. Acta Zoologica, DOI: 10.1111/j.1463-6395.2011.00502.x

Abstract. The microstructure of dorsal osteoderms referred to the chroniosuchid taxa Chroniosuchus, Chroniosaurus, Madygenerpeton and cf. Uralerpeton is compared to existing data on the bystrowianid chroniosuchian Bystrowiella and further tetrapods. Chroniosuchid osteoderms are marked by thin internal and relatively thick external cortices that consist of lowly vascularised parallel-fibred bone. They are structured by growth marks and, in case of Madygenerpeton, by lines of arrested growth. The cancellous middle region is marked by a high degree of remodelling and a primary bone matrix of parallel-fibred bone that may include domains of interwoven structural fibres. Whereas the convergence of Bystrowiella and chroniosuchid osteoderms is not confirmed by our observations, the internal cortex of the latter displays a significant peculiarity: It contains distinct bundles of shallowly dipping Sharpey’s fibres with a cranio- or caudoventral orientation. We interpret this feature as indicative for the attachment of epaxial muscles which spanned several vertebral segments between the medioventral surface of the osteoderms and the transversal processes of the thoracic vertebrae. This finding endorses the hypothesis that the chroniosuchid osteoderm series was part of a crocodylian-like trunk bracing system that supported terrestrial locomotion. According to the measured range of osteoderm bone compactness, some chroniosuchian species may have had a more aquatic lifestyle than others.

Three recent papers on chroniosuchians

Buchwitz M, Voigt S. 2010. Peculiar carapace structure of a Triassic chroniosuchian implies evolutionary shift in trunk flexibility. Journal of Vertebrate Paleontology 30: 1697-1708. [Link]

Schoch RR, Voigt S, Buchwitz M. 2010. A chroniosuchid from the Triassic of Kyrgyzstan and analysis of chroniosuchian relationships. Zoological Journal of the Linnaean Society 160: 515-530. [Link]

Klembara J, Clack J, Čerňanský A. 2010. The anatomy of palate of Chroniosaurus dongusensis (Chroniosuchia, Chroniosuchidae) from the Upper Permian of Russia. Palaeontology 53: 1147-1153. [Link]

The redescription of the Chroniosaurus dongusensis palate by Klembara and colleagues adds further data to the morphological dataset provided by Clack and Klembara (2009) in their revision of C. dongusensis on the basis of a new specimen (which is the most complete of any yet known chroniosuchian). According to the updated phylogenetic analysis from the 2010 paper Chroniosaurus as the only included chroniosuchian taxon formed the sister group of embolomeres.

Schoch and colleagues (me included) describe Madygenerpeton pustulatus, a new species of chroniosuchians from the Middle to Late Triassic of Central Asia with a highly derived skull morphology and a carapace that was chroniosuchid-like in many aspects. The find shows that one lineage of chroniosuchids survived the Permian-Triassic boundary (by 20 or so million years).

The authors discuss characteristics uniting chroniosuchians with "higher reptiliomorphs" and unlike the approach of Klembara and colleagues their cladistic analysis, which includes five chroniosuchian taxa, results in a position of chroniosuchians somewhat closer to amniotes than to embolomeres. Chroniosaurus comes out as the closest relative of Madygenerpeton (both share the characteristic ornamentation of the skull and osteoderms besides other features).

Buchwitz & Voigt consider the functionality of chroniosuchian carapaces, comparing them to archosaur osteoderm systems. They argue that chroniosuchian carapaces basically served terrestrial locomotion but that the higher lateral flexibility of the Madygenerpeton osteoderm system was linked to a secondary increase in undulation swimming capability.

Reference:
Clack JA, Klembara J. 2009. An articulated specimen of Chroniosaurus dongusensis, and the morphology and relationships of the chroniosuchids. Special Papers in Palaeontology 81: 15-42. [Link]

Reptile from the Petrified Forest of Chemnitz

In Early Permian volcaniclastic deposits of Chemnitz (Saxony, Germany) the partial skeleton of 30-cm-long basal reptile including a complete autopodium, further parts of the limbs, the vertebral column, thorax, and a fragmentary skull have been discovered last week by excavators from the Natural History Museum of Chemnitz.

This is the first tetrapod fossil from these deposits which are otherwise famous for their in situ silified tree stems ("Petrified Forest of Chemnitz").

Chroniosuchians and stay in Moscow

To resolve the riddle: The bonified eyeball from the last post represents a a ball-shaped intercentrum of a chroniosuchid from the Permian of Russia.

In chroniosuchian reptiliomorphs the intercentra (white arrows) are interlocked with the amphicoelous pleurocentra in a ball-and-socket-like fashion. The image on the left shows some section of a Chroniosuchus vertebral column in ventral view. Intercentra become bony balls only in the adult individuals - not fully bonified in sub-adults and juveniles they are preserved with a crescent, disk-like or ellipsoidal shape.

Also, the fusion of the neural arch with the pleurocentrum - a feature otherwise characteristic for "higher reptiliomorphs" such as seymouriamorphs and diadectomorphs - is only completed in the course of ontogenesis (so that you can find suture lines in the subadult individuals).

Moscow: Paleontological Institute and Museum of the Russian Academy of Sciences (PIN)

The paleo-style PIN building has a castle-like rectangular shape with an inner courtyard featuring life-size sculptures of fossil critters from Russia and areas of the former Soviet Union. The exhibition has almost everything you wish for as a vertebrate enthusiast (here depicted: the two-story dinosaur hall). Rich in type specimens the collection is essential for some and important for many studies - so earlier or later many of the fossil vertebrate people spend some time there.

Concering the Chroniosuchia: With the exception of bystrowianid chroniosuchian remains from Kupferzell, Germany (Witzmann et al. 2008) and China (Young 1979) and some rather questionable Chinese chroniosuchid chroniosuchians (Li & Cheng 1999), all yet described Permian and Triassic chroniosuchian taxa come from the European part of Russia and are mostly archived in the PIN. I am thankful to Valery Golubev who helped me a lot during my week of stay when I was studying the type materials and to Jury Gubin who nicely put up with me in his room.

Some literature on chroniosuchians

Here considered: titles also available in English (plus the Chinese ones mentioned above).

Golubev, V. K. (1998). "Narrow-armored Chroniosuchians (Amphibia, Anthracosauromorpha) from the Late Permian of Eastern Europe." Paleontologicheskij Zhurnal 1998(3): 64- 73. [Russian, English]

Golubev, V. K. (1998). "Revision of the Late Permian chroniosuchians (Amphibia, Anthracosauromorpha) from Eastern Europe." Paleontologicheskij Zhurnal 1998(4): 68- 77. [Russian, English]

Golubev, V. K. (1999). "A new narrow-armored chroniosuchian (Amphibia, Anthracosauromorpha) from the Late Permian of the East Europe." Paleontologicheskij Zhurnal 1999(2): 43- 50. [Russian, English]

Li, J., Cheng Z. (1999). " New anthracosaur and temnospondyl amphibians from Gansu, China." Vertebrata PalAsiatica 37(3): 234- 247. [Chinese with English abstract]

Novikov, I. V., M.A. Shishkin (2000). "Triassic chroniosuchians (Amphibia, Anthracosauromorpha) and the evolution of the trunk dermal ossifications in the bystrowianids." Paleontological Journal 34(supplement): S165- S178. [English]

Novikov, I. V., M.A. Shishkin, V.K. Golubev (2000). Permian and Triassic anthracosaurs from Eastern Europe. The Age of Dinosaurs in Russia and Mongolia. M. A. S. M.J. Benton, D.M. Unwin, E.N. Kurochkin. Cambridge, Cambridge University Press: 60- 70. [English]

Witzmann, F., R.R. Schoch, M.W. Maisch (2008). "A relic basal tetrapod from the Middle Triassic of Germany." Naturwissenschaften 95(1): 67- 72. [English]

Young, C. C. (1979). "A new Late Permian fauna from Jiyuan, Honan." Vertebrata Palasiatica 17: 99- 113. [Chinese with English abstract]

Bonified eyeball

Or what the hell is this?