Madygen Formation

Coordinates: 40°06′N 70°12′E / 40.1°N 70.2°E / 40.1; 70.2
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Madygen Formation
Stratigraphic range: Carnian
~235–222 Ma
TypeGeological formation
OverliesCambrian to Carboniferous rocks
Thickness560 m (1,840 ft)
Lithology
PrimaryConglomerate, sandstone
OtherMudstone
Location
Coordinates40°06′N 70°12′E / 40.1°N 70.2°E / 40.1; 70.2
Approximate paleocoordinates41°12′N 60°36′E / 41.2°N 60.6°E / 41.2; 60.6
RegionBatken & Osh Regions
CountryKyrgyzstan
Tajikistan
Uzbekistan
ExtentFergana Valley & Range
Type section
Named forMadygen village
Named byEvgeny A. Kochnev
Madygen Formation is located in Kyrgyzstan
Madygen Formation
Madygen Formation (Kyrgyzstan)

The Madygen Formation (Russian: Madygen Svita) is a Late Triassic (Carnian) geologic formation and Lagerstätte in the Batken and Osh Regions of western Kyrgyzstan, with minor outcrops in neighboring Tajikistan and Uzbekistan. The conglomerates, sandstones and mudstones of the 560 m (1,840 ft) thick formation were deposited in terrestrial lacustrine, alluvial, fluvial and deltaic environments.

The formation, extending across the Fergana Valley and Fergana Range, is unique for Central Asia, as it represents one of the few known continental deposits and the Madygen Formation is renowned for the preservation of more than 20,000 fossil insects, making it one of the richest Triassic Lagerstätten in the world. Other vertebrate fossils as fish, amphibians, reptiles and synapsids have been recovered from the formation too, as well as minor fossil flora.

The lake sediments of the Lagerstätte provided fossil cartilaginous fishes and their egg capsules and unusual Triassic reptiles like Sharovipteryx and Longisquama.[1][2] The wide diversity of insect fossils was first discovered in the 1960s and first described by Russian paleontologist Aleksandr Sharov, with a notable example being Gigatitan.[3]

Description[edit]

Paleogeography of the Late Triassic, around 230 Ma. The Madygen Formation was deposited north of the Paleo-Tethys ocean.

The Madygen Formation is a 560 metres (1,840 ft)[4] thick succession of predominantly siliciclastic rocks accumulated in a tectonically induced basin, covering parts of the Fergana Range and Fergana Valley of Kyrgyzstan with minor outcrops in Tajikistan and Uzbekistan.[5] The Late Triassic layers rest on top of Paleozoic basement with local Permo-Triassic molasse sediments. The section consists of mudstones, sandstones, conglomerates and fanglomerates. This wide variety of siliciclastic rocks reflects the complex spatial and temporal pattern of depositional sub-environments including alluvial fans, sandflats, swamps, back-swamp areas, and littoral to profundal lake zones. The fluvio-lacustrine deposits of the Madygen Formation belong to one of only a few occurrences of continental Triassic beds in Central Asia.[6]

The formation was deposited during the Carnian Pluvial Event (CPE), a global humid event leading to high extinction levels of various groups globally. The CPE led to anoxic conditions, most notably in the South China Block. The area where the Madygen Formation was deposited formed part of the Cimmerian microcontinent, a slab of crust that collided with Laurasia during the Cimmerian orogeny in later Mesozoic times. This orogeny led to the disappearance of the Paleo-Tethys Ocean.

Petroleum geology[edit]

The formation grades from bottom to top from alluvial to fluvial into a thick succession of lacustrine mudstones, followed by an alluvial package, on top of which lacustrine, fluvial, deltaic and alluvial layers were deposited.[5]

The hydrocarbon potential of samples of the Madygen Formation ranges from poor to excellent. The sediments containing more than 0.5% Total Organic Carbon (TOC) may be regarded as sources of gaseous hydrocarbons rather than of oil.[5] The Hydrogen Index (HI) of outcrop samples reaches 100 and the maximum recorded maturity (Ro) is 0.8.[5]

Paleontological significance[edit]

During the 1960s, Russian paleontologists recovered an unusually rich fossil content in the type strata of the Madygen Formation, including abundant macrophytes, more than 20,000 insect remains[7] and unique small reptiles with well preserved soft tissue.[6] Spirorbis-like polychaete worm tubes, crustaceans (ostracods, kazacharthrans, malacostraca), freshwater Bivalves and gastropods are known from shallow to deeper lake environments. Non-aquatic insects are among the most common fossil remains of the Madygen Formation. These include representatives of the orders Ephemeroptera, Odonata, Notoptera, Blattodea, Titanoptera, Ensifera, Caelifera, Rhynchota, Auchenorrhyncha, Stenorrhyncha, Coleoptera, Hymenoptera, Trichoptera and Diptera. Traces of insect larvae are preserved in near-shore lake deposits.[8]

Fish remains mostly represent endemic genera assigned to the actinopterygian families Evenkiidae (Oshia), Palaeoniscidae (Ferganiscus, Sixtelia) and Megaperleidus and Alvinia. The actinopterygian Saurichthys and the dipnoan Asiatoceratodus are cosmopolitan taxa also recorded in the Madygen Formation. Two distinctive elasmobranch egg capsule types, i.e. Palaeoxyris, indicating a small Lissodus- or Lonchidion-like hybodont shark and an indeterminate capsule type, imply the presence of two different elasmobranch species which used the freshwater environments of the Madygen Formation as spawning grounds. Tetrapods are known from a probably larval urodelan (Triassurus), a small procynosuchid cynodont (Madysaurus), a gliding reptile (Sharovipteryx) and the enigmatic diapsid Longisquama.[8]

Paleobiota[edit]

Color key
Taxon Reclassified taxon Taxon falsely reported as present Dubious taxon or junior synonym Ichnotaxon Ootaxon Morphotaxon
Notes
Uncertain or tentative taxa are in small text; crossed out taxa are discredited.

Amphibians[edit]

Genus Species Material Notes Images
Madygenerpeton[9] M. pustulatus[9] Skull and osteoderms. A chroniosuchid reptiliomorph with aquatic adaptations.[9][10]
Triassurus T. sixtelae plentiful remains, mostly of larval individuals The earliest known member of the caudata (salamanders and their kin). Most of the known fossils include larval remains (the neural arches of the vertebrae were still paired and no vertebral centers show any degree of ossification).[11]

Reptiles[edit]

Genus Species Material Notes Images
Kyrgyzsaurus[12] K. bukhanchenkoi[12] A single specimen preserving the front half of a skeleton and scale impressions.[12] One of, if not the oldest member of the drepanosaurs, a diverse grouping of Triassic reptiles that evolved arboreal, fossorial, and potentially aquatic lifestyles.[13] It was also the first Asian member of the group to be described.[12]
Longisquama[14] L. insignis[14] A specimen preserving the front half of a skeleton and "plumes", and at least five additional "plume" fragments.[2] A genus of neodiapsid reptile that is notable due to the holotype specimen possessing large growths on its back. This animal has gone through a confusing taxonomic history, with some authors suggesting a placement within the archosauromorpha, and as a basal diapsid.[14][2][15]
Sharovipteryx S. mirabilis[16] A single skeleton with impressions of gliding membranes, split across a slab and counterslab.[2] A gliding archosauromorph, and a member of the sharovipterygidae family. Originally named as Podopteryx,[16] a genus name which was preoccupied by a damselfly.[17][2]

Synapsids[edit]

Genus Species Material Notes Images
Madysaurus[18] M. sharovi[18] A single partial skeleton. A cynodont[18]

Cartilaginous fishes[edit]

Possible xenacanth denticles, egg cases, as well as hybodont fossils have also been reported from the formation.[19]

Genus Species Material Notes Images
Fayolia F. sharovi[1] Egg capsules An egg capsule likely belonging to a xenacanthid.[1] The egg is elongate and tapers towards both ends, and surrounded by helically twisted collarettes, with one end (the beak) having a tendril.[20]
Lonchidion L. ferganensis[1] Teeth, denticles,[19] and egg capsules A hybodontid fish that lived from the Lower Triassic to the Upper Cretaceous. The fossils found at Madygen (which mainly come from juvenile individuals) suggest this animal spawned within freshwater areas.[1]
Palaeoxyris P. alterna[1] Teeth and egg capsules The egg cases of hybodonts that appear frequently throughout the fossil record.[1] They comprise a beak, a body and a pedicle. They display a conspicuous right-handed spiral of collarettes around the body, and in some cases, the pedicle, resulting in a rhomboidal pattern when flattened during fossilisation.

Bony fishes[edit]

The following fish fossils were found in the formation:[21][8][22][19]

Genus Species Material Notes Images
Alvinia[21] A. serrata[21] Partial skeletons. A small perleidid.[8][22]
Asiaceratodus A. sharovi Partial skeletons. A medium-sized dipnoan (lungfish), around 30 centimetres (12 in) in length.[8][22]
Ferganiscus[21] F. osteolepis[21] Nearly complete skeletons. A small and abundant palaeoniscid.[8][22]
Megaperleidus[21] M. lissolepis[21] Partial skeletons. A medium-sized perleidid.[8][22]
Oshia[21] O. ferganica[21] Partial skeletons, scales.[19] A medium-sized evenkiid scanilepiform with predatory habits. Around 45 centimetres (18 in) in length.[22]
Saurichthys S. orientalis[21] Nearly complete skeleton,[22] scales,[19] other fragmentary material. A medium-sized saurichthyid chondrostean, around 45 centimetres (18 in) in length.[22]
Sixtelia[21] S. asiatica[21] Nearly complete skeletons, scales.[19] A small and abundant palaeoniscid.[8][22]

Arthropods[edit]

Genus Species Description Notes Images
Gigatitan G. extensus A mantis-like titanopteran insect with a wingspan of approximately 40 centimetres (16 in).[23] It is the type genus of the family Gigatitanidae. Characteristics of the wings (they were able to produce flashes) suggest this insect was a diurnal predator.[24] [25]
G. magnificus
G. ovatus
G. similis
G. vulgaris
Aiban A. kichineis A member of Cnemidolestida/Cnemidolestodea (an extinct group of insects of uncertain phylogenetic placement, might be related to plecopterans or orthopterans) belonging to the family Sylvabestiidae [26]
Batkentak B. intactus A member of Grylloblattida/Eoblattida belonging to the family Daldubidae [27]
Chubakka C. madygensis A madygelline xyelid sawfly [28]
Locustoblattina L. marginata A member of Eoblattida belonging to the family Mesorthopteridae [29]
L. segmentata
Madygella M. aristovi A madygelline xyelid sawfly [28]
M. bashkuevi
M. kurochkini
M. levivenosa
Nestorembia N. novojilovi A webspinner belonging to the family Alexarasniidae [30]
N. shcherbakovi [29]
Paratitan P. reliquia A right wing of an insect originally identified as a late-surviving palaeodictyopteran (Liquia reliquia)[31] but subsequently reinterpreted as a titanopteran.[32] [31]
Sharovites S. alexanderi A member of Grylloblattida/Eoblattida, belonging to the family Mesorthopteridae [33]
Shurabia S. tanga A member of Polyneoptera belonging to the group Reculida and the family Geinitziidae [34]
Guillermia G. lecticula [35]
Madygenorhynchus M. multifidus
Thaumatomerope T. sogdiana
Prochoristella P. longa
Dilemmala D. specula
Sacvoyagea S. ventrosa
Nonescyta N. mala
Maguviopsis M. kotchnevi
Falcarta F. bella
Krendelia K. ansata
Cuanoma C. protracta
Phyllotexta P. latens
Sitechka S. perforata
Tingiopsis T. reticulata
Pelorisca P. connectens
Xamenophlebia X. ornata
Thuringoblatta T. sogdianensis
Sacvoyagea S. ventrosa
Protoblattogryllus P. variabilis
Megablattogryllus M. austerus
M. pinguis
Metakhosara M. sharovi
Batkentak B. intactus
Austroidelia A. asiatica
A. nervosa
Mesoidelia M. faceta
Parastenaropodites P. fluxa
P. longiuscula
Baharellus B. madygensis
Anoblattogryllus A. fundatus
Costatoviblatta C. aenigmatosa
C. conjuncta
Paratitan P. reductus
Prototitan P. sharovi
Mesoedischia M. obliqua
Paragryllavus P. curvatus
Zagryllavus Z. elongatus
Hagloedischia H. primitiva
Voliopus V. ancestralis
Euvoliopus E. giganteus
Macrovoliopus M. declivis
Paravoliopus P. dorsalis
Voliopellus V. latus
Triassaga T. angusta
Zamaraga Z. reticulata
Haglomorpha H. martynovi
Modihagla M. ovalis
Dulcihagla D. mistshenkoi
Lyrohagla L. uvarovi
Sonohagla S. curta
Tinnihagla T. zeuneri
Dolichohagla D. longa
Locustavus L. problematicus
L. intermedius
L. minutus
Miolocustavus M. reductus
Brevilocustavus B. distinctus
Fritaniopsis F. brevicaulis
Sogdoblatta S. nana
S. porrecta
Thuringoblatta T. sogdianensis
Xiphopterum X. sharovi
Sharovoplana S. parallelica
Triassophasma T. intermedium
T. pusillum
Prochresmoda P. parva
Nestorembia N. novojilovi
Madygenophlebia M. bella
M. nana
Gorochovia G. individua
G. minuta
G. bifurca
Gorochoviella G. conjuncta
Pseudoliomopterites P. obscurus
Ideliopsina I. nana
I. stupenda
I. ornata
Pseudoshurabia P. pallidula
Peltosyne P. varyvrosa
Ofthalmopeltos O. synkritos
Obrienia O. illaetabilis
Triassochorista T. kirgizica
Parachorista P. arguta
P. immota
Choristopanorpa C. temperata
Thaumatomerope T. oligoneura
T. sogdiana
Mesageta M. rieki
Liassochorista L. utilis
Agetopanorpa A. deceptoria
Prochoristella P. longa
Mesopsyche M. ordinata
M. justa
M. tortiva
Psychotipa P. predicta
Vymrhyphus V. tuomikoskii
Gnomusca G. molecula
Asiocula A. lima
Fasolinka F. beckermigdisovae
Fulgobole F. evansi
Scytachile S. emeljanovi
Serpentivena S. tigrina
Coccavus C. supercubitus
Kennedya K. carpenteri
K. gracilis
Batkenia B. pusilla
Paurophlebia P. lepida
P. angusta
Neritophlebia N. elegans
N. vicina
N. longa
Mixophlebia M. mixta
Cyrtophlebia C. sinuosa
Zygophlebia Z. ramosa
Reisia R. sogdiana
Shurabia S. serrata [36]
Mesoblattogryllus M. intermedius [25]
Protoblattogryllus P. asiaticus
P. variabilis
Megablattogryllus M. austerus
M. magister
Megakhosarodes M. paulivenosus
Metakhosara M. sharovi
Austroidelia A. asiatica
A. nervosa
Mesoidelia M. ignorata
M. faceta
Parastenaropodites P. longiuscula
P. fluxa
Locustoblattina L. segmentata
Dorniella D. primitiva
Baharellus B. madygensis
Baharellinus B. dimidiatus
B. pectinatus
Costatoviblatta C. aenigmatosa
C. similis
Ferganamadygenia F. plicata
Paratitan P. libelluloides
P. venosus
P. intermedius
P. latispeculum
P. modestus
Mesotitanodes M. tillyardi
M. similis
Mesotitan M. primitivus
Provitimia P. pectinata
Proshiella P. ramivenosa
Mesoedischia M. madygenica
Gryllacrimima G. perfecta
Madygenia M. orientalis
M. ovalis
Kashgarlimahmutia K. reducta
Proxenopterum P. primitivum
Axenopterum A. venosum
Ferganopterus F. clarus
Ferganopterodes F. reductus
Pteroferganodes P. rieki
Tuphella T. rohdendorfi
T. sharovi
Platyvoliopus P. maximus
Stenovoliopus S. elongatus
Zavoliopus Z. densus
Turkestania T. deviata
Triassaga T. tshorkuphlebioides
Eumaraga E. madygenica
Hagloptera H. intermedia
Archihagla A. tenuis
Proisfaroptera P. martynovi
Protshorkuphlebia P. triassica
P. similis
Dulcihagla D. beybienkoi
Lyrohagla L. pravdini
L. decipiens
Sonohagla S. saussurei
S. chopardi
Microhagla M. minuta
Dinohagla D. corrugata
Adzhajloutshella A. plana
Locustavus L. lanceolatus
L. problematicus
L. intermedius
L. deformatus
L. minutus
Dicronemoura D. acaulis
Tritaniella T. mera
T. synneura
Fritaniopsis F. brevicaulis
F. remota
Sogdoblatta S. nana
Thuringoblatta T. sogdianensis
Subioblatta S. madygenica
Xiphopterum X. curvatum
Sharovoplana S. affinis
Triassophasma T. brevipoda
T. intermedium
T. minutissimum
Prochresmoda P. minuta
Madygenophlebia M. bella
M. primitiva
Micromadygenophlebia M. obscura
Gorochovia G. individua
Ideliopsina I. nana
I. ornata
Madygenidelia M. conjuncta
Pseudoshurabia P. pallidula
Hadeocoleus H. gigas
H. pelopius
H. catachtonius
Triassocoleus T. tortulosus
Salebroferus S. confragosus
S. asper
Schizophoroides S. rugosus
Thnesidius T. xyphophorus
Catabrycus C. hoplites
Triaplus T. laticoxa
Dolichosyne D. rostrata
Ademosyne A. kirghizica
Cephalosyne C. capitata
Cupesia C. sepulta
Notocupes N. laticella
N. rostratus
N. tenuis
Cladochorista C. multivenosa
Prophilopotamus P. asiaticus
Triassochorista T. kirgizica
Parachorista P. asiatica
P. multivena
P. arguta
P. sana
Mesageta M. pertrita
M. ignava
Agetopanorpa A. consueta
Prochoristella P. longa
Mesopsyche M. shcherbakovi
M. justa
M. gentica
Psychotipa P. depicta
Nadiptera N. pulchella
Oryctoxyela O. anomala
Madygenius M. primitivus
Ferganoxyela F. sogdiana
F. destructa
Triassoxyela T. orycta
Nevicia N. imitans
Asiocula A. lima
Phyllotexta P. latens
Fulgobole F. evansi
Scytachile S. cf. emeljanovi
Tingiopsis T. reticulata
Coccavus C. supercubitus
Permonka P. unica
P. triassica
Permosialis P. triassica
Triassolestodes T. asiaticus
Terskeja T. pumilio
T. tenuis
Paurophlebia P. lepida
Cladophlebia C. parvula
C. brevis
Nonymophlebia N. venosa
Neritophlebia N. longa
Triadophlebia T. madygenica
T. minuta
T. magna
T. honesta
T. modica
Mitophlebia M. enormis
Xamenophlebia X. ornata
Zygophlebiella Z. curta
Mixophlebia M. mixta
Shurabia S. minuta
S. ferganensis
S. anomala
Maguviopsis M. kotchnevi [37]
Tingiopsis T. reticulata
Dolichosyne D. confragosa [38]
Rhabdocupes R. baculatus
Salebroferus S. confragosus
Asiocula A. lima [39]
Blattomerope B. polyneura
Cuanoma C. protracta
Fasolinka F. beckermigdisovae
Kirgizichorista K. larvata
Krendelia K. ansata
Lithocupes L. incertus
Maguviopsis M. kotchnevi
Megakhosarodes M. paulivenosus
Nonescyta N. mala
Obrienia O. ingurgata
O. kuscheli
Panorpaenigma P. aemulum
Parachorista P. religiosa
Phyllotexta P. latens
Sacvoyagea S. ventrosa
Siberioperla S. ovalis
Sitechka S. perforata
Sogdodromeus S. altus
Triaplus T. macroplatus
Megakhosarodes M. paulivenosus [40]
Choristopanorpa C. opinata [41]
Dzhajloutshella D. arcanum
Kuperwoodia K. benefica
Macrocatinius M. brachycephalus
Neotuphella N. minor
Pesus P. prognathus
Mesageta M. gigantea [42]
M. rieki [41]
Triassoxya T. novozhilovi [43]

Flora[edit]

The plant fossils of the Madygen Formation were reviewed in detail by Dobruskina (1995).[44]

Genus Species Description Notes Images
Mesenteriophyllum M. kotschnevii [15]

(partial list)

Insect fauna correlations[edit]

Progonocimicidae found in the formation are also recorded in the Carnian Los Rastros Formation of Argentina, the Norian Blackstone and Mount Crosby Formations of Australia, and the Norian to Rhaetian Tologoi Formation of Kazakhstan.[7] Permochoristidae are also known from the Carnian Potrerillos and Cacheuta Formations of Argentina, Huangshanjie Formation of China, the Norian Blackstone and Mount Crosby Formations of Australia; the Norian to Rhaetian Tologoi Formation of Kazakhstan, the Sinemurian Dzhil Formation of Kyrgyzstan and the Toarcian Posidonia Shale of Germany.[7]

Orthophlebia had a relatively broad distribution in the Late Triassic as it is also found in the Sinemurian Badaowan Formation of China and Dzhil Formation of Kyrgyzstan, the Pliensbachian Makarova Formation of Russia and Sulyukta Formation of Tajikistan; the Toarcian Whitby Mudstone Formation of England, Posidonia Shale of Germany, and Cheremkhovo Formation of Russia, and the Early Jurassic Kushmurun Formation of Kazakhstan.[7]

Haglidae were also recorded in the Koldzat and Tologoi Formations of Kazakhstan, in the Carnian Cacheutá Formation of Argentina, the Carnian to Norian Molteno Formation of South Africa and Lesotho, and the Norian Mount Crosby Formation of Australia.[7]

See also[edit]

Other Central Asian Lagerstätten

References[edit]

  1. ^ a b c d e f g Fischer, Jan; Voigt, Sebastian; Schneider, Jörg W.; Buchwitz, Michael; Voigt, Silke (2011). "A selachian freshwater fauna from the Triassic of Kyrgyzstan and its implication for Mesozoic shark nurseries". Journal of Vertebrate Paleontology. 31 (5): 937–953. doi:10.1080/02724634.2011.601729. ISSN 0272-4634.
  2. ^ a b c d e Unwin, David M.; Alifanov, Vladimir R.; Benton, Michael J. (2003). "Enigmatic small reptiles from the Middle-Late Triassic of Kirgizstan". In Benton, Michael J.; Shishkin, Mikhail A.; Unwin, David M.; Kurochkin, Evgenii M. (eds.). The Age of Dinosaurs in Russia and Mongolia. Cambridge: Cambridge University Press. pp. 177–186. ISBN 9780521545822.
  3. ^ Shcherbakov, Dmitry (2008). "Madygen, Triassic Lagerstätte number one, before and after Sharov" (PDF). Alavesia. 2: 113–124.
  4. ^ Fischer, Jan; Kogan, Ilja; Voigt, Sebastian; Buchwitz, Michael; Schneider, Jörg W.; Moisan, Philippe; Spindler, Frederik; Brosig, Andreas; Preusse, Marvin; Scholze, Frank; Linnermann, Ulf (2018). "The mid-Triassic Madygen Lagerstätte (Southwest Kyrgyzstan, Central Asia)". 13th Symposium on Mesozoic Terrestrial Ecosystems and Biota. Bonn, Germany: 25–26.
  5. ^ a b c d Berner, Ulrich; Scheeder, Georg; Kus, Jolanta; Voigt, Sebastian (2009). "Organic Geochemical Characterization of Terrestrial Source Rocks of the Triassic Madygen Formation (Southern Tien Shan, Kyrgyzstan)". OIL GAS European Magazine. 35 (3): 135–139.
  6. ^ a b Fischer, Jan; Voigt, Sebastian; Buchwitz, Michael (2007). "First elasmobranch egg capsules from freshwater lake deposits of the Madygen Formation (Middle to Late Triassic, Kyrgyzstan, Central Asia)". Freiberger Forschungshefte (C 524): 41–46.
  7. ^ a b c d e Richard S., Kelly (2018). "Effects of environmental perturbation during the Late Triassic on the taxic diversity of British insects" (PDF). University of Bristol Earth Sciences PhD dissertation: 1–337.
  8. ^ a b c d e f g h Voigt, Sebastian; Spindler, Frederik; Fischer, Jan; Kogan, Ilja; Buchwitz, Michael (2007). "An extraordinary lake basin – the Madygen fossil lagerstaette (Middle to Upper Triassic, Kyrgyzstan, Central Asia)". Wissenschaftliche Mitteilungen des Institutes für Geologie der TU Bergakademie Freiberg. 36: 162–163.
  9. ^ a b c Schoch, Rainer R.; Voigt, Sebastian; Buchwitz, Michael (2010). "A chroniosuchid from the Triassic of Kyrgyzstan and analysis of chroniosuchian relationships". Zoological Journal of the Linnean Society. 160 (3): 515–530. doi:10.1111/j.1096-3642.2009.00613.x.
  10. ^ Buchwitz, Michael; Voigt, Sebastian (2010-12-02). "Peculiar carapace structure of a Triassic chroniosuchian implies evolutionary shift in trunk flexibility". Journal of Vertebrate Paleontology. 30 (6): 1697–1708. doi:10.1080/02724634.2010.521685. ISSN 0272-4634.
  11. ^ Schoch, Rainer R.; Werneburg, Ralf; Voigt, Sebastian (2020-05-26). "A Triassic stem-salamander from Kyrgyzstan and the origin of salamanders". Proceedings of the National Academy of Sciences. 117 (21): 11584–11588. doi:10.1073/pnas.2001424117. ISSN 0027-8424. PMC 7261083. PMID 32393623.
  12. ^ a b c d Alifanov, V. R.; Kurochkin, E. N. (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. ISSN 0031-0301.
  13. ^ Jenkins, X.A.; Pritchard, A.C.; Marsh, A.D.; Kligman, B.T.; Sidor, C.A.; Reed, K.E. (2020-10-08). "Using manual ungual morphology to predict substrate use in the Drepanosauromorpha and the description of a new species". Journal of Vertebrate Paleontology. 40 (5): e1810058. doi:10.1080/02724634.2020.1810058. ISSN 0272-4634. S2CID 225136171.
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  23. ^ Park, Tae-Yoon S.; Kim, Do-Yoon; Nam, Gi-Soo; Lee, Mirinae (2022-05-06). "A new titanopteran Magnatitan jongheoni n. gen. n. sp. from southwestern Korean Peninsula". Journal of Paleontology. 96 (5): 1111–1118. doi:10.1017/jpa.2022.30. ISSN 0022-3360. S2CID 248592776.
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  25. ^ a b Sharov, 1968
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  30. ^ Shcherbakov, D. E. (2015). "Permian and Triassic ancestors of webspinners (Embiodea)". rej (in Russian). 24 (1): 187–200. doi:10.15298/rusentj.24.3.01. ISSN 0132-8069.
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  33. ^ Aristov, D.S.; Storozhenko, S. Yu. (2013). "A new genus of the family Mesorthopteridae (Grylloblattida) from the Triassic of Kyrgyzstan". Far Eastern Entomologist. 264: 7–12.
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  37. ^ Becker-Migdisova, 1953
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  39. ^ Arnoldi et al., 1977
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Further reading[edit]

  • Gorochov, A.V. 2005. Review of Triassic Orthoptera with Descriptions of New and Little Known Taxa: Part 2. Paleontological Journal 39(3). 272–279. .
  • Novokshonov, V.G. 1997. New Triassic Scorpionflies (Insecta, Mecoptera). Paleontological Journal 31. 628–635. .
  • Shcherbakov, D.E.; E.D. Lukashevich, and V.A. Blagoderov. 1995. Triassic Diptera and initial radiation of the order. International Journal of Dipterological Research 6. 75-115. .
  • Storozhenko, S.Y. 1993. Reviziya semeystva Megakhosaridae (Grylloblattida) In A. G. Ponomarenko (ed.). Mezozoyskie Nasekomye i Ostrakody Azii .. 100–112. .
  • Arnoldi, L.V.; V.V. Zherikhin; L.M. Nikritin, and A.G. Ponomarenko. 1977. Mezozoiskie zhestkokryiye. Akademiya Nauk SSSR, Trudy Paleontologicheskogo Instituta 161. 1–204. .
  • Ponomarenko, A.G. 1969. Istoricheskoe Razvitie Zhestkokrylykh-Arkhostemat [Historical Development of the Archostomate Beetles]. Trudy Akademiya Nauk SSSR 125. 1–240. .
  • Sharov, A.G. 1968. Filogeniya ortopteroidnykh nasekomykh. Trudy Paleontologicheskogo Instituta Akademii Nauk SSSR 118. 1–216. .
  • Becker-Migdisova, E.E. 1953. Dva predstavitelya poluzhestkokrylykh nasekomykh iz urochishcha Madygen. Doklady Akademii Nauk SSSR 90. 461–464. .
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