Madygen Formation

The Madygen Formation (Russian: Madygen Svita) is a Middle–Late Triassic (Ladinian–Carnian) geologic formation and lagerstätte in the Batken and Osh Regions of western Kyrgyzstan. It also has small outcrops in nearby Tajikistan and Uzbekistan. This place is important because it has many fossils from long ago, showing what life was like millions of years ago.

The Madygen Formation is well-known for having more than 20,000 fossil insects. This makes it one of the richest places in the world for finding fossils from the Triassic period. Scientists have also found fossils of fish, amphibians, reptiles, and other early animals here. These fossils help us learn about the ancient worlds and ecosystems that existed there.

The lake sediments in the Madygen Formation have given us amazing fossils, including early cartilaginous fishes and their eggs. They also have unusual reptiles like Sharovipteryx and Longisquama. The many insect fossils were first found in the 1960s by Russian paleontologist Aleksandr Sharov, and one famous example is Gigatitan. This formation helps scientists understand life from over 200 million years ago.

Description

The Madygen Formation is a thick layer of rocks that is about 560 metres (1,840 ft) tall. It is found in parts of the Fergana Range and Fergana Valley in Kyrgyzstan, with some pieces also in Tajikistan and Uzbekistan. These rocks are made from materials like mud, sand, and small rounded stones.

These rocks tell us about many different places where they were formed long ago, such as areas near rivers, swamps, and lakes. They are special because they are one of the few places in Central Asia with rocks from the time of the dinosaurs that were not under the sea. Scientists have found many fossils in these rocks, especially in an area called Dzaylyaucho, which looks like it was once a big, deep lake far from any ocean.

Paleontological significance

In the 1960s, scientists found many important fossils in the Madygen Formation. They discovered over 20,000 fossils of insects, along with small reptiles. Fossils of plants, worms, tiny sea creatures, and freshwater shells were also found in the lake areas.

The Madygen Formation is special because it has many insect fossils. These insects belong to many different groups. The fossils also include fish and early reptiles, showing the variety of life that lived there long ago.

Paleobiota

Amphibians

Reptiles

Reptiles are the most common group of four-legged animals at Madygen, with three described types.

Synapsids

Cartilaginous fishes

Possible xenacanth tooth-like structures, egg cases, and hybodont fossils have been found in the formation. The presence of many young individuals suggests that the freshwater systems at Madygen served as spawning grounds and nurseries for these ancient elasmobranchs.

Bony fishes

The following fish fossils were found in the formation:

Insects

The Madygen Formation is famous for its insect fossils, with many specimens and types found in the formation. Some believe it has the most diverse insect group from the Permian-Triassic time period.

Odonata (damselflies, dragonflies and kin)

The Odonata and other Odonatoptera of the Madygen Formation are rare, with moderate variety.

• †Paurophlebiidae (8 types): Cladophlebia (C. brevis, C. parvula), Neritophlebia (N. elegans, N. longa, N. vicina), Nonymophlebia (N. venosa), Paurophlebia (P. angusta, P. lepida)
• †Triadophlebiidae (6 types): Triadophlebia (T. distincta, T. honesta, T. madygenica, T. magna, T. minuta, T. modica)
• †Zygophlebiidae (4 types): Cyrtophlebia (C. sinuosa), Mixophlebia (M. mixta), Zygophlebia (Z. ramosa), Zygophlebiella (Z. curta)
• †Kennedyidae (4 types): Kennedya (K. carpenteri, K. ferganensis, K. gracilis, K. madygensis)
• †Voltzialestidae (3 types): Terskeja (T. paula, T. pumilio, T. tenuis)
• †Batkeniidae (2 types): Batkenia (B. pusilla), Paratriassoneura (P. primitiva)
• †Mitophlebiidae (1 type): Mitophlebia (M. enormis)
• †Protomyrmeleontidae (1 type): Ferganagrion (F. kirghiziensis)
• †Triadotypidae (1 type): Reisia (R. sogdiana)
• †Triassolestidae (1 type): Triassolestodes (T. asiaticus)
• †Xamenophlebiidae (1 type): Xamenophlebia (X. ornata)

Blattodea (roaches)

The Blattodea of the Madygen Formation are very common, and some show details of the whole body. Roaches in the families †Caloblattinidae and †Spiloblattinidae are especially common and diverse.

• †Caloblattinidae: Sogdoblatta (S. maxima, S. nana, S. porrecta), Thuringoblatta (T. sogdianensis)
• †Subioblattidae: Subioblatta (S. madygenica)
• Undescribed types of †Blattulidae, †Phyloblattidae, †Poroblattinidae, †Spiloblattinidae, and potentially †Archimylacridae.

†Titanoptera

The †Titanoptera of the Madygen Formation are uncommon, with moderate variety. One titanopteran of special note is Gigatitan vulgaris, a mantis-like diurnal hunter with a wingspan of approximately 40 centimetres (16 in).

• †Paratitanidae (11 types): Microtitan (M. zherichini), Paratitan (P. bispeculum, P. intermedius, P. latispeculum, P. libelluloides, P. longispeculum, P. modestus, P. ovalis, P. reductus, P. reliquia, P. venosus)
• †Mesotitanidae / †Gigatitanidae (11 types): Gigatitan (G. ovatus, G. similis, G. vulgaris), Mesotitanodes (M. tillyardi), Nanotitan (N. extentus, N. magnificus), Ootitan (O. curtis), Prototitan (P. primitivus, P. sharovi, P. similis), Ultratitan (U. superior)

Note: Béthoux (2007) considers †Titanoptera to simply be a subgroup of orthopterans, rather than its own group.

Orthoptera (grasshoppers and kin)

The Orthoptera of the Madygen Formation are very common and very diverse. Most types belong to the subgroup Ensifera (relatives of crickets, katydids, and grigs), with only a few Caelifera (relatives of grasshoppers). None of the families survive to today, and most are only found in the Triassic.

• †Haglidae (44 types): Ahagla (A. mira), Archihagla (A. tenuis, A. zeuneri), Cantohagla (C. gracilis), Dinohagla (D. corrugata), Dolichohagla (D. longa), Dulchihagla (D. beybienkoi, D. mistshenkoi), Eumaraga (E. madygenica), Euvoliopus (E. giganteus), Haglomorpha (H. martynovi), Hagloptera (H. intermedia), Lyrohagla (L. decipiens, L. pravdini, L. uvarovi), Macrovoliopus (M. declivis), Maragella (M. reducta), Melovoliopus (M. fasciatus), Microhagla (M. minuta), Modihagla (M. ovalis), Paravoliopus (P. dorsalis), Phonovoliopus (P. musicus), Platyvoliopus (P. maximus), Proisfaroptera (P. martynovi), Protshorkuphlebia (P. kirgizica, P. similis, P. triassica), Sharovohagla (S. plana), Sonohagla (S. chopardi, S. curta, S. saussurei), Stenovoliopus (S. elongatus), Tinnihagla (T. handlirschi, T. zeuneri), Triassaga (T. angusta, T. tshorkuphlebioides), Turkestania (T. deviata), Vocohagla (V. clara, V. tarbinskyi), Voliopellus (V. latus), Voliopus (V. ancestralis), Zamaraga (Z. recticulata), Zavoliopus (Z. densus), Zeunerophlebia (Z. gigas)
• †Proparagryllacrididae (16 types): Batkenella (B. megaptera), Brevibatkenella (B. abscisa), Dolichobatkenella (D. perlonga), Eubatkenella (E. devexia), Gryllacrimima (G. elongata, G. madygenioides, G. perfecta, G. simplicis), Kashgarlimahmutia (K. reducta), Madygenia (M. extremalis, M. longissima, M. orientalis, M. ovalis), Oedischimima (O. deficientis), Parafergania (P. sharovi), Platymadygenia (P. grandis)
• †Xenopteridae (15 types): Axenopterum (A. venosum), Ferganopterodes (F. reductus), Ferganopterus (F. clarus, F. longus), Ferganotriassia (F. lata), Proxenopterum (P. primitivum), Pseudoferganopsis (P. zini), Pteroferganella (P. crassa, P. minuta, P. sharovi), Pteroferganodes (P. decipiens, P. rieki), Triassoferganella (T. angusta), Triassomanteodes (T. madygenicus), Xenoferganella (X. pini)
• †Locustavidae (9 types): Brevilocustavus (B. distinctus, B. microscopicus), Ferganopsis (F. lanceolatus), Locustavus (L. deformatus, L. intermedius, L. madygensis, L. minutus, L. problematicus), Miolocustavus (M. reductus)
• †Bintoniellidae (8 types): Oshiella (O. crassa, O. oblonga), Oshiellana (O. primaria), Paroshiella (P. alia), Probintoniella (P. triassica), Proshiella (P. ramivenosa), Provitimia (P. pectinata), Stenoshiella (S. angusta)
• †Dzhajloutshellidae (5 types): Adzhajloutshella (A. planis, A. talis), Dzhajloutshella (D. arcanum, D. flexuosa), Triassoxya (T. novozhilovi)
• †Tuphellidae (4 types): Neotuphella (N. minor), Tuphella (T. rasnitsyni, T. rohdendorfi, T. sharovi)
• †Gryllavidae (3 types): Gryllavus (G. madygenicus), Paragryllavus (P. curvatus), Zaragryllavus (Z. elongatus)
• †Mesoedischiidae (3 types): Mesoedischia (M. kirgizia, M. madygenica, M. obliqua)
• †Permelcanidae (2 types): Meselcana (M. madygenica, M. permelcanoides)
• †Hagoedischiidae (1 type): Hagoedischia (H. primitiva)

Phasmatodea (stick insects and kin)

The Phasmatodea of the Madygen Formation are somewhat common and somewhat low in variety. All of the families were only found in the Triassic.

• †Prochresmodidae (8 types): Prochresmoda (P. longipoda, P. media, P. minuta, P. parva), Triassophasma (T. brevipoda, T. intermedium, T. minutissimum, T. pusillum)
• †Aeronautidae (2 types): Sharovoplana (S. affinis, S. parallelica)
• †Xiphopteridae? (2 types): Xiphopterum (X. curvatum, X. sharovi)

Grylloblattodea (ice-crawlers and kin) and other basic polyneopterans

The Grylloblattodea and other basic polyneopterans of the Madygen Formation (sometimes described as Eoblattida or Protorthoptera, among other names) are very common and diverse.

• †Megakhosaridae (11 types): Madygenocephalus (M. micropteron), Megablattogryllus (M. austerus, M. magister, M. pinguis), Megakhosarodes (M. obtusus, M. paulovenosus), Mesoblattogryllus (M. abruptus, M. intermedius), Metakhosara (M. sharovi), Protoblattogryllus (P. asiaticus, P. variabilis)
• †Mesorthopteridae (11 types): Belmophenopterum (B. rasnitsyni), Austroidelia (A. asiatica), Locustoblattina (L. marginata, L. segmentata), Mesoidelia (M. faceta, M. rasnitsyni, M. semota), Parastenaropodites (P. fluxa, P. longiuscula, P. nervosa), Sharovites (S. alexanderi_)_
• †Blattogryllidae (9 types): Anoblattogryllus (A. fundatus), Baharellinus (B. dimidiatus, B. pectinatus), Baharellus (B. madygensis), Costatoviblatta (C. aenigmatosa, C. conjuncta, C. longipennis, C. similis_), Dorniella (D. primitiva)_
• †Gorochoviidae (8 types): Gorochovia (G. anomala, G. bifurca, G. fecunda, G. individua, G. minuta), Gorochoviella (G. conjuncta), Pseudoliomopterites (P. lucidus, P. obscurus)
• †Ideliidae (7 types): Anaidelia (A. extrema), Ideliopsina (I. nana, I. ornata, I. ruginosa, I. stupenda), Madygenidelia (M. conjuncta), Pseudoshurabia (P. pallidula)
• †Geinitziidae (5 types): Geinitziella (G. rasnitsyni_)_, Shurabia (S. anomala, S. ferganensis, S. minutissima, S. tanga)
• †Madygenophlebiidae (4 types): Madygenophlebia (M. bella, M. nana, M. primitiva), Micromadygenophlebia (M. obscura)
• †Sylvabestiidae (2 types): Aiban (A. kichineis_)_, Sharovala (S. triassica)
• †Necrophasmatidae (1 type): Ferganamadygenia (F. plicata)
• †Daldubidae (1 type): Batkentak (B. intactus)
• †Sylvaphlebiidae (1 type): Batkenopterum (B. kirgizicum_)_

†Miomoptera

The †Miomoptera of the Madygen Formation are somewhat common, most or all of which belong to a single valid type.

• †Permosialidae (1 type): Permosialis (P. triassica_)_

Plecoptera (stoneflies)

The Plecoptera of the Madygen Formation are rare and somewhat low in variety.

• †Perlariopseidae (14 types): Cristonemoura (C. binerva, C. porrecta), Dicronemoura (D. acaulis, D. declinata, D. dira), Fritaniopsis (F. brevicaulis, F. dependens, F. remota), Ramonemoura (R. constricta), Triassonemoura (T. ficteramosa), Tritaniella (T. mera, T. pectinata, T. perlonga, T. synneura)
• †Siberioperlidae (1 type): Siberioperla (S. ovalis)

Embioptera (webspinners)

The Embioptera of the Madygen Formation are rare and very low in variety.

• †Alexarasniidae: Nestorembesia (N. novojilovi, N. shcherbakovi)
• †Rasnalexiidae: Rasnalexia (R. rasnitsyni)

Hemiptera (true bugs)

The Hemiptera of the Madygen Formation are very common and very diverse, though most remain undescribed. By far the most common are small cicadomorphs (relatives of treehoppers, leafhoppers, and cicadas), though large butterfly-like cicadas also occur.

• †Maguvopseidae: Asiocula (A. lima), Cuanoma (C. protracta), Falcarta (F. bella), Fasolinka (F. beckermigdisovae), Krendelia (K. ansata), Maguviopsis (M. kotchnevi), Nevicia (N. imitans), Nonescyta (N. mala), Phyllotexta (P. latens), Sacvoyagea (S. ventrosa), Sitechka (S. perforata)
• †Palaeontinidae: Papiliontina (P. dracomima, P. machaon, P. spectans)
• †Mesojabloniidae: Fulgobole (F. evansi), Mesojablonia (M. kukalovae), Scytachile (S. emeljanovi)
• †Dunstaniidae: Dunstaniodes (D. elongatus), Siksteliana (S. popovi)
• †Curvicubitidae: Beaconiella (B. cincta, B. pulchra)
• †Creaphididae: Creaphis (C. theodora)
• †Ipsviciidae: Strivicia (S. davidi)
• †Naibiidae: Coccavus (C. supercubitus_)_
• †Paraknightiidae: Triknightia (T. mira)
• †Progonocimicidae: Pelorisca (P. connectens)
• †Saaloscytinidae: Tingiopsis (T. reticulata)
• †Serpentivenidae: Serpentivena (S.tigrina)
• Undescribed types of †Chiliocyclidae, †Dysmorphoptilidae, †Hylicellidae, †Mesogereonidae, †Pincombeidae, †Protopsyllidiidae, †Scytinopteridae, †Stenoviciidae, †Surijocixiidae, and possibly Ochteroidea.

Neuroptera (lacewings and kin)

The Neuroptera of the Madygen Formation are uncommon but diverse in form. Only a few types have been formally described.

• †Archeosmylidae: Madygoneura (M. elongata), Osmylotriasia (O. superba), Triasella (T. ovata)
• Berothidae (beaded lacewings): Ferganoberotha (F. miniutissima)
• †Permithonidae: Relictovia (R. pristina)
• Undescribed types of †Osmylopsychopidae, Osmylidae (lance lacewings), and "Polystoechotidae" (giant lacewings).

Coleoptera (beetles)

The Coleoptera of the Madygen Formation are very common and diverse. By far the most common beetles are archostematans, particularly †Schizophoridae and cupedid-grade beetles (†Triadocupedidae, Ommatidae).

• †Schizophoridae (17 types): Catabrycus (C. hoplites), Hadeocoleus (H. catachtonius, H. gigas, H. pelopius), Lethocoleus (L. sternalis), Pesus (P. prognathus), Praesagus (P. capitatus), Salebroferus (S. asper, S. confragosus), Schizophorinus (S. punctatus), Schizophoroides (S. glaber, S. rugosus, S. tuberculatus), Thnesidius (T. ovatus, T. xyphophorus), Triassocoleus (T. sulcatus, T. tortulosus)
• †Triadocupedidae / †Triadocupedinae (16 types): Asimma (A. rara), Cupesia (C. monilicornia, C. sepulta, C. serricornia), Kirghizocupes (K. cellulosus), Platycupes (P. dolichocerus, P. major, P. pusillus, P. reticulatus, P. sogdianus), Procupes (P. mandibularis), Pterocupes (P. antennatus, P. leptocerus), Triadocupes (T. ellipticus, T. ferghanensis, T. latus)
• Ommatidae / Ommatinae (reticulated beetles, 12 types): Lithocupes (L. gigas, L. incertus, L. punctatus), Notocupes? (N. laticella, N. rostratus, N. tenuis), Notocupoides? (N. capitatus, N. fasciatus, N. triassicus), Rhabdocupes? (R. baculatus, R. longus, R. minor)
• †Ademosynidae (7 types): Ademosyne (A. bacca, A. elliptica, A. kirghizica), Cephalosyne (C. capitata), Dolichosyne (D. confragosa, D. rostrata, D. sulcata)
• †Triaplidae (7 types): Avocatinus (A. elongatus), Catinoides (C. rotundatus), Macrocatinius (M. brachycephalus), Triaplus (T. laticoxa, T. macroplatus), Triassocatinius (T. brachynotus, T. glabratus)
• †Obrieniidae (5 types): Guillermia (G. lecticula), Madygenorhynchus (M. multifidus), Obrienia (O. illaetabilis, O. ingurgata, O. kuscheli)
• †Peltosynidae (4 types): Gnathopeltos (G. dixis), Ofthalmopeltos (O. synkritos), Peltosyne (P. triassica, P. varyvrosa)
• †Asiocoleidae / †Tricoleidae (3 types): Sogdelytron (S. latum), Tricoleodus (T. acutus, T. longus)
• Cupedidae sensu stricto / Cupedinae (reticulated beetles, 2 types): Mesocupoides (M. indistinctus, M. proporeius)
• Trachypachidae (false ground beetles, 1 type): Sogdodromus (S. altus)

Hymenoptera (wasps and kin)

The Hymenoptera of the Madygen Formation are rare, but with moderate variety. All the types belong to the sawfly family Xyelidae, which still survives to today.

• Xyelidae (xyelid sawflies, 36 types): Asioxyela (A. parvula, A. paurura, A. smilodon), Chubakka (C. madygensis), Dinoxyela (D. armata), Euryxyela (E. euryptera, E. lata), Ferganoxyela (F. destructa, F. sogdiana), Leioxyela (L. antiqua, L. grandis, L. kirgizica, L. mitis, L. mollis), Lithoxyela (L. fenestralis), Madygella (M. analoga, M. aristovi, M. bashkuevi, M. kurochkini, M. levivenosa), Madygenius (M. extraradius, M. primitivus), Oryctoxyela (O. anomala, O. triassica), Microxyela (M. minuta), Samarkandykia (S. ryzhkovae, S. shmakovi_), Triassoxyela (T. foviolata, T. grandipennis, T. orycta, T. sharovi), Xiphoxyela (X. procrusta, X. striata), Xyelinus (X. angustiradius, X. major, X. scherbachov)_

Mecoptera (scorpionflies)

The Mecoptera of the Madygen Formation are very common and fairly diverse.

• †Permochoristidae (15 types): Agetopanorpa (A. consueta, A. deceptoria, A. triassica), Liassochorista (L. molesta, L. utilis), Mecolusor (M. confusicus), Mesageta (M. gigantea, M. ignava, M. insana, M. pertrita, M. rieki), Mesochorista (M. injuriosa), Prochoristella (P. ignara, P. longa, P. vicina)
• †Parachoristidae (13 types): Choristopanorpa (C. opinata, C. ridibunda, C. temperata), Kirgizichorista (K. larvata), Panorpaenigma (P. aemulum), Parachorista (P. arguta, P. asiatica, P. comica, P. immota, P. multivena, P. religiosa, P. sana), Triassochorista (T. kirgizica)
• †Thaumatomeropidae (6 types): Blattomerope (B. polyneura), Pronotiothauma (P. neuropteroides), Thaumatomerope (T. madygenica, T. minuta, T. oligoneura, T. sogdiana)
• †Mesopsychidae (5 types): Mesopsyche (M. gentica, M. justa, M. ordinata, M. shcherbakovi, M. tortiva)
• incertae sedis (2 types): †Mecaenigma (M. suspectum, M. viduum)
• †Pseudopolycentropodidae (1 type): Pseudopolycentropus (P. madygenicus)

Trichoptera (caddisflies)

The Trichoptera of the Madygen Formation are rare and low in variety.

• †Cladochoristidae: Cladochorista (C. curta), Cladochoristella (C. sola)
• †Prorhyacophilidae: Prorhyacophila (P. batkenica, P. furcata, P. rara,)
• †Necrotauliidae: Paranecrotaulius (P. proximus)
• Philopotamidae? Prophilopotamus (P. asiaticus) (dubious)

Diptera (flies)

The Diptera of the Madygen Formation are rare and somewhat low in variety.

• †Psychotipidae (2 types): Psychotipa (P. depicta, P. predicta)
• †Protorhyphidae (2 types): Vymrhyphus (V. triassicus, V. tuomikoskii)
• †Gnomuscidae? (2 types): Gnomusca (G. molecula, G. renyxa)
• Chaoboridae (phantom midges, 1 type): Triassomyia (T. shcherbakovi_)_
• †Hennigmatidae? (1 type): Anemeca (A. liya)
• †Kuperwoodidae (1 type): Kuperwoodia (K. benefica)
• Limoniidae (limoniid crane flies, 1 type): Mabelysia (M. charlesi)
• †Nadipteridae (1 type): Nadiptera (N. pulchella)
• †Vladipteridae (1 type): Dilemmala (D. specula)

Other insects

Psocoptera (book lice) are represented by a single undescribed type from the family †Psocidiidae. Dermaptera (earwigs) are also very rare and undescribed, represented by the family †Protodiplateidae. The extinct order †Glosselytrodea is only slightly more diverse, with rare fossils from the families †Jurinidae and †Polycytellidae. The only fossil Ephemeroptera (mayfly) reported from the formation is a Mesobaetis-like fragment, and its referral to a mayfly has been doubted.

Other invertebrates

Apart from insects, the most common invertebrates found in the Madygen Formation are Almatium gusevi and Jeanrogerium sornayi, tadpole shrimp-like aquatic crustaceans in the order Kazacharthra. Undescribed ostracod and decapod fossils have also been found, though conchostracans are practically absent. The Madygen Formation is one of the only Triassic sites to preserve statoblasts of freshwater bryozoans (Phylactolaemata). Freshwater bivalves, gastropods, microconchid shells, and worm burrows are also found in the lakebed sediments.

Flora

The plant fossils of the Madygen Formation were reviewed in detail by Dobruskina (1995). Among the oldest known root nodules have been found in the Madygen Formation, though the exact nature of the plant-microbe interaction responsible remains unknown.

Non-vascular plants

Ferns, horsetails, and lycopods

Ferns are rare and poorly-preserved, though certain horsetails (Neocalamites) and lycopods (Ferganadendron, Mesenteriophyllum, Isoetites) are more common.

Pteridosperms ("seed ferns")

Pteridosperms are abundant and diverse, making up more than half of all the plant fossils recovered from the formation.

Other gymnosperms

Though subordinate to pteridosperms in terms of abundance, other gymnosperms are still quite diverse, and some of them (Glossophyllum, Podozamites, Taeniopteris) are far from uncommon.

Insect fauna correlations

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. 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.

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.

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.

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