Madygen Formation
The Madygen Formation is a Middle–Late Triassic 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 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. 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.
Description
The Madygen Formation is a 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. The Middle-Late Triassic layers rest on top of Paleozoic basement with local Permo-Triassic molasse sediments. They are likewise overlain by latest-Triassic to Early Jurassic sediments, some of which host their own arthropod faunas.The formation consists of a wide and multicolored variety of siliciclastic rocks: mudstones, sandstones, conglomerates, fanglomerates, and rare coal layers. This diversity of sediment types reflects the a complex set of depositional environments through time and space, 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.
Isotope analyses of fish teeth confirm that the lakes and rivers represented by the formation were entirely freshwater and deep inland, about 600 km from the nearest coast. Though fossils can be found throughout the entire formation, the most extensive fossil deposits occur in the northern area, known as Dzaylyaucho, which is dominated by brown and grey claystones and mudstones. The abundance of burrows and absence of darker sediments in this area indicate that it was a well-oxygenated lacustrine environment, such as a large oxbow lake.
The environment represented by the Madygen Formation was positioned on 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. The Madygen Formation was emplaced at warm temperate latitudes in a rift basin surrounded by old, eroded mountains, the ancestral Tien Shan Mountains. The climate was likely humid and moist year-round, as drying features such as mudcracks and evaporite deposits are absent.
Petroleum geology
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.The hydrocarbon potential of samples of the Madygen Formation ranges from poor to excellent. The sediments containing more than 0.5% Total Organic Carbon may be regarded as sources of gaseous hydrocarbons rather than of oil. The Hydrogen Index of outcrop samples reaches 100 and the maximum recorded maturity is 0.8.
Paleontological significance
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 and unique small reptiles with well preserved soft tissue. Spirorbis-like polychaete worm tubes, crustaceans, 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, with half a thousand or so species having been uncovered from these sediments. This makes the Madygen Formation one of the most prolific insect fossil sites In the world. These include representatives of both extinct and extant orders, including the 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.Fish remains mostly represent endemic genera assigned to the actinopterygian families Evenkiidae, Palaeoniscidae 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 the mostly larval urodelan, a small procynosuchid cynodont, a chroniosuchid reptiliomorph, an early drepanosaurid reptile, a gliding archosauromorph and the enigmatic diapsid Longisquama.
Paleobiota
Reptiles
Reptiles are the most abundant of the tetrapod paleofauna at Madygen, with three described genera.| Genus | Species | Material | Notes | Images |
| Kyrgyzsaurus | K. bukhanchenkoi | A single specimen preserving the front half of a skeleton and scale impressions. | Potentially the oldest known member of the drepanosaurs, a diverse clade of Triassic reptiles that evolved arboreal and fossorial lifestyles. It was also the first member of the group to be described from Asia. | |
| Longisquama | L. insignis | A specimen preserving the front half of a skeleton and "plumes", and at least five additional "plume" fragments. | A neodiapsid reptile possessing unusual large scaly growths on its back. This animal has gone through a confusing taxonomic history, with some authors suggesting a placement within Archosauromorpha, or as a basal diapsid. | |
| Sharovipteryx | S. mirabilis | A single skeleton with impressions of gliding membranes, split across a slab and counterslab. | A gliding archosauromorph, and the type genus of the family Sharovipterygidae. Originally named as Podopteryx, a genus name which was preoccupied by a damselfly. |
Cartilaginous fishes
Possible xenacanth denticles, egg cases, as well as hybodont fossils have also been reported from the formation. The high amount of juvenile individuals found suggest the freshwater systems at the Madygen Formation served as a spawning grounds and nursery's for these prehistoric elasmobranchs.| Genus | Species | Material | Notes | Images |
| Fayolia | F. sharovi | Egg capsules | An egg capsule likely belonging to a xenacanthid. The egg is elongate and tapers towards both ends, and surrounded by helically twisted collarettes, with one end having a tendril. | |
| Palaeoxyris | P. alterna | Egg capsules | The egg cases of hybodonts that appear frequently throughout the fossil record. 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. The eggs were most likely produced by the contemporary genus Lonchidion, due to the number of preserved juvenile individuals. | |
| Lonchidion | L. ferganensis | Teeth, denticles, and egg capsules | A hybodontid elasmobranch that lived from the Lower Triassic to the Upper Cretaceous. The fossils found at Madygen suggest this animal spawned within freshwater areas. The majority of the teeth from the Madygen Formation originated from areas full of abundant bivalve fossils, suggesting the young hybodonts frequented these areas due to the rich food supply. |
Bony fishes
The following fish fossils were found in the formation:| Genus | Species | Material | Notes | Images |
| Alvinia | A. serrata | Partial skeletons. | A small perleidid. | |
| Asiatoceratodus | A. sharovi | Partial skeletons. | A medium-sized dipnoan, around in length. | |
| Ferganiscus | F. osteolepis | Nearly complete skeletons. | A small and abundant palaeoniscid, in length. | |
| Megaperleidus | M. lissolepis | Partial skeletons. | A medium-sized perleidid. | |
| Oshia | O. ferganica | Partial skeletons, scales. | A medium-sized evenkiid scanilepiform with predatory habits. Around in length. | |
| Saurichthys | S. orientalis | Nearly complete skeleton, scales, other fragmentary material. | A relatively small saurichthyid chondrostean, around in length. | |
| Sixtelia | S. asiatica | Nearly complete skeletons, scales. | A small and common palaeoniscid, in length. |
Insects
The Madygen Formation is renowned for its insect fauna, with an estimated 25,000 specimens, 500 species, 100 families, and 20 orders found in the formation. Under some estimates, it has the most diverse insect assemblage of the entire Permian-Triassic interval. From, Shcherbakov, and the Paleobiology Database, unless cited otherwise:Odonata (damselflies, dragonflies and kin)
The Odonata and other Odonatoptera of the Madyen Formation are rare, with moderate diversity. They were described most extensively by Pritykina.- †Paurophlebiidae : Cladophlebia, Neritophlebia, Nonymophlebia, Paurophlebia
- †Triadophlebiidae : Triadophlebia
- †Zygophlebiidae : Cyrtophlebia, Mixophlebia, Zygophlebia, Zygophlebiella
- †Kennedyidae : Kennedya
- †Voltzialestidae : Terskeja
- †Batkeniidae : Batkenia, Paratriassoneura
- †Mitophlebiidae : Mitophlebia
- †Protomyrmeleontidae : Ferganagrion
- †Triadotypidae : Reisia
- †Triassolestidae : Triassolestodes
- †Xamenophlebiidae : ''Xamenophlebia ''
Blattodea (roaches)
The Blattodea of the Madygen Formation are extremely abundant, and some preserve details of the entire body. Roaches in the families †Caloblattinidae and †Spiloblattinidae are particularly common and diverse, though most species remain undescribed. Some species were given a preliminary description by Vishniakova.- †Caloblattinidae: Sogdoblatta, Thuringoblatta
- †Subioblattidae: Subioblatta
- Undescribed species of †Blattulidae, †Phyloblattidae, †Poroblattinidae, †Spiloblattinidae, and potentially †Archimylacridae.
†Titanoptera
The †Titanoptera of the Madygen Formation are uncommon, with moderate diversity, though still higher than any other geological formation. They were described most extensively by Sharov and Gorochov. One titanopteran of special note is Gigatitan vulgaris, one of the few insects from the formation known from more than wings. This species was a superficially mantis-like diurnal predator with a wingspan of approximately 40 centimetres. Like other large titanopterans, it was probably a poor flier which primarily used its wings to produce sounds or flashes of light for courtship of defensive purposes.- †Paratitanidae : Microtitan, Paratitan
- †Mesotitanidae / †Gigatitanidae : Gigatitan, Mesotitanodes, Nanotitan, Ootitan, Prototitan, Ultratitan
Orthoptera (grasshoppers and kin)
The Orthoptera of the Madygen Formation are very common and very diverse. Most species belong to the suborder Ensifera, with only a few Caelifera. None of the families survive to the present, and most are endemic to the Triassic. Madygen orthopterans were described most extensively by Sharov and Gorochov.- †Haglidae : Ahagla, Archihagla, Cantohagla, Dinohagla, Dolichohagla, Dulchihagla, Eumaraga, Euvoliopus, Haglomorpha, Hagloptera, Lyrohagla, Macrovoliopus, Maragella, Melovoliopus, Microhagla, Modihagla, Paravoliopus, Phonovoliopus, Platyvoliopus, Proisfaroptera, Protshorkuphlebia, Sharovohagla, Sonohagla, Stenovoliopus, Tinnihagla, Triassaga, Turkestania, Vocohagla, Voliopellus, Voliopus, Zamaraga, Zavoliopus, Zeunerophlebia
- †Proparagryllacrididae : Batkenella, Brevibatkenella, Dolichobatkenella, Eubatkenella, Gryllacrimima, Kashgarlimahmutia, Madygenia, Oedischimima, Parafergania, Platymadygenia
- †Xenopteridae : Axenopterum, Ferganopterodes, Ferganopterus, Ferganotriassia, Proxenopterum, Pseudoferganopsis, Pteroferganella, Pteroferganodes, Triassoferganella, Triassomanteodes, Xenoferganella
- †Locustavidae : Brevilocustavus, Ferganopsis, Locustavus, Miolocustavus
- †Bintoniellidae : Oshiella, Oshiellana, Paroshiella, Probintoniella, Proshiella, Provitimia, Stenoshiella
- †Dzhajloutshellidae : Adzhajloutshella, Dzhajloutshella, Triassoxya
- †Tuphellidae : Neotuphella, Tuphella
- †Gryllavidae : Gryllavus, Paragryllavus, Zaragryllavus
- †Mesoedischiidae : Mesoedischia
- †Permelcanidae : Meselcana
- †Hagloedischiidae : ''Hagloedischia ''
Phasmatodea (stick insects and kin)
The Phasmatodea of the Madygen Formation are somewhat common and somewhat low in diversity. All of the families were endemic to the Triassic. They were described most extensively by Sharov and Gorochov.- †Prochresmodidae : Prochresmoda, Triassophasma
- †Aeroplanidae : Sharovoplana
- †Xiphopteridae? : ''Xiphopterum ''
Grylloblattodea (ice-crawlers and kin) and other basal polyneopterans
The Grylloblattodea and other basal polyneopterans of the Madygen Formation are very common and diverse. They were described most extensively by Storozhenko.- †Megakhosaridae : Madygenocephalus, Megablattogryllus, Megakhosarodes, Mesoblattogryllus, Metakhosara, Protoblattogryllus
- †Mesorthopteridae : Belmophenopterum, Austroidelia, Locustoblattina, Mesoidelia, Parastenaropodites, Sharovites
- †Blattogryllidae : Anoblattogryllus, Baharellinus, Baharellus, Costatoviblatta, Dorniella
- †Gorochoviidae : Gorochovia, Gorochoviella, Pseudoliomopterites
- †Ideliidae : Anaidelia, Ideliopsina, Madygenidelia, Pseudoshurabia
- †Geinitziidae : Geinitziella , Shurabia
- †Madygenophlebiidae : Madygenophlebia, Micromadygenophlebia
- †Sylvabestiidae : Aiban , Sharovala
- †Necrophasmatidae : Ferganamadygenia ''''
- †Daldubidae : Batkentak
- †Sylvaphlebiidae : ''Batkenopterum ''
†Miomoptera
The †Miomoptera of the Madygen Formation are somewhat common, most or all of which belong to a single valid species.- †Permosialidae : ''Permosialis ''
Plecoptera (stoneflies)
The Plecoptera of the Madygen Formation are rare and somewhat low in diversity. They were described most extensively by Sinitchenkova.- †Perlariopseidae : Cristonemoura, Dicronemoura, Fritaniopsis, Ramonemoura, Triassonemoura, Tritaniella
- †Siberioperlidae : ''Siberioperla ''
Embioptera (webspinners)
The Embioptera of the Madygen Formation are rare and very low in diversity. Some Madygen mesorthopterids show similarities to alexarasniids, which may suggest that †Mesorthopteridae is ancestral to the embiopteran lineage.- †Alexarasniidae: Nestorembesia
- †Rasnalexiidae: Rasnalexia
Hemiptera (true bugs)
The Hemiptera of the Madygen Formation are extremely abundant and very diverse, though most remain undescribed. By far the most common are small cicadomorphs, though large butterfly-like cicadas also occur. Fulguroidea and Stenorrhyncha are present but uncommon. The only heteropteran in the formation is an extremely rare basal Nepomorpha similar to Ochteroidea.- †Maguvopseidae: Asiocula, Cuanoma, Falcarta, Fasolinka, Krendelia, Maguviopsis, Nevicia, Nonescyta, Phyllotexta, Sacvoyagea, Sitechka
- †Palaeontinidae: Papiliontina
- †Mesojabloniidae: Fulgobole, Mesojablonia, Scytachile
- †Dunstaniidae: Dunstaniodes, Siksteliana
- †Curvicubitidae: Beaconiella
- †Creaphididae: Creaphis
- †Ipsviciidae: Strivicia
- †Naibiidae: Coccavus
- †Paraknightiidae: Triknightia
- †Progonocimicidae: Pelorisca
- †Saaloscytinidae: Tingiopsis
- †Serpentivenidae: Serpentivena
- Undescribed species 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 species have been formally described.- †Archeosmylidae: Madygoneura, Osmylotriasia, Triasella
- Berothidae : Ferganoberotha
- †Permithonidae: Relictovia
- Undescribed species of †Osmylopsychopidae, Osmylidae, and "Polystoechotidae".
Coleoptera (beetles)
The Coleoptera of the Madygen Formation are extremely abundant and diverse. By far the most abundant beetles are archostematans, particularly †Schizophoridae and cupedid-grade beetles. They were described most extensively by Ponomarkenko.- †Schizophoridae : Catabrycus, Hadeocoleus, Lethocoleus, Pesus, Praesagus, Salebroferus, Schizophorinus, Schizophoroides, Thnesidius, Triassocoleus
- †Triadocupedidae / †Triadocupedinae : Asimma, Cupesia, Kirghizocupes, Platycupes, Procupes, Pterocupes, Triadocupes
- Ommatidae / Ommatinae : Lithocupes, Notocupes?, Notocupoides?, Rhabdocupes?
- †Ademosynidae : Ademosyne, Cephalosyne, Dolichosyne
- †Triaplidae : Avocatinus, ''Catinoides, Macrocatinius, Triaplus, Triassocatinius
- †Obrieniidae : Guillermia, Madygenorhynchus, Obrienia
- †Peltosynidae : Gnathopeltos, Ofthalmopeltos, Peltosyne
- †Asiocoleidae / †Tricoleidae : Sogdelytron, Tricoleodus
- Cupedidae sensu stricto / Cupedinae : Mesocupoides
- Trachypachidae : Sogdodromus ''
Hymenoptera (wasps and kin)
The Hymenoptera of the Madygen Formation are rare, but with moderate diversity. All the species belong to the sawfly family Xyelidae, which still survives to the present. They were described most extensively by Rasnitsyn.- Xyelidae : ''Asioxyela, Chubakka, Dinoxyela, Euryxyela, Ferganoxyela, Leioxyela, Lithoxyela, Madygella, Madygenius, Oryctoxyela, Microxyela, Samarkandykia, Triassoxyela, Xiphoxyela, Xyelinus ''
Mecoptera (scorpionflies)
The Mecoptera of the Madygen Formation are very common and fairly diverse. They were described most extensively by Ponomarenko & Rasnitsyn, and Novokshonov.- †Permochoristidae : Agetopanorpa, Liassochorista, Mecolusor, Mesageta, Mesochorista, Prochoristella
- †Parachoristidae : Choristopanorpa, Kirgizichorista, ''Panorpaenigma, Parachorista, Triassochorista
- †Thaumatomeropidae : Blattomerope, Pronotiothauma, Thaumatomerope
- †Mesopsychidae : Mesopsyche incertae sedis : †Mecaenigma
- †Pseudopolycentropodidae : Pseudopolycentropus ''
Trichoptera (caddisflies)
The Trichoptera of the Madygen Formation are rare and low in diversity.- †Cladochoristidae: Cladochorista, Cladochoristella
- †Prorhyacophilidae: Prorhyacophila
- †Necrotauliidae: Paranecrotaulius
- Philopotamidae? ''Prophilopotamus ''
Diptera (flies)
The Diptera of the Madygen Formation are rare and somewhat low in diversity. They were described most extensively by Shcherbakov et al.. Some purported Madygen dipterans wings may belong to four-winged insects instead.- †Psychotipidae : Psychotipa
- †Protorhyphidae : Vymrhyphus
- †Gnomuscidae? : Gnomusca
- Chaoboridae : Triassomyia
- †Hennigmatidae? : Anemeca
- †Kuperwoodidae : Kuperwoodia
- Limoniidae : Mabelysia
- †Nadipteridae : Nadiptera
- †Vladipteridae : ''Dilemmala ''
Other insects
Psocoptera are represented by a single undescribed species from the family †Psocidiidae. Dermaptera 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 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. 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. 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.Ferns, horsetails, and lycopods
Ferns are rare and poorly-preserved, though certain horsetails and lycopods are more common.| Genus | Species | Notes | Images |
| Chiropteris | C. integalla | Fern fronds | |
| Cladophlebis | C. ex. gr. nebbensis | Osmundacean fern fronds | |
| Cladophlebis | C. ex. gr. paralobifolia | Osmundacean fern fronds | |
| Cladophlebis | C. raciborskii | Osmundacean fern fronds | |
| Cladophlebis | C. ex. gr. shensiensis | Osmundacean fern fronds | |
| Ctenopteris | C. punctata | A dubious fern-like plant of uncertain affinities | |
| Danaeopsis | D. fecunda | Marattiacean fern fronds | |
| Equisetites | E. sp. | Stems and other fragments of a small sphenophyte | |
| Ferganadendron | F. sauktangensia | Bark, stem fragments, and tiny attached leaves of a lycopsid. Sometimes described as a tree-like plant with a stem up to in diameter, but most fossils suggest a much smaller maximum diameter of. Previously misidentified as a species of Sigillaria. | |
| Isoetites | I. madygensis | Quillwort corms with attached leaves, very similar to modern Isoetes. Also probably includes lycopod scales previously named as Annalepis leae. Both Madygen Isoetites species occasionally exhibit rows of deformed scars, likely produced via the egg-laying behavior of damselfly-like insects in the extinct suborder Archizygoptera. | |
| Isoetites | I. sixteliae | Serrated quillwort leaves, quite common. | |
| Mesenteriophyllum | M. kotschnevii | Enigmatic strap-like leaves, up to long, with a pleated texture and in some cases serrated margins. Most likely from a type of lycopsid. Previously considered to be two species, M. kotschnevii and M. serratum, with the former characterized by large spines, and the latter characterized by finer serrations. It is notable due to a study from 2006 presenting the possibility that the plumes of Longisquama were potentially plant leaves that had gotten preserved alongside the diapsid. However, Buchwitz & Voigt rebuked this claim, stating that the plumes do not resemble the shape of this plant's leaves, and that they are not preserved in carbon films. | |
| Neocalamites | N. hoerensis | Leafy stems and other fragments of a large and very common sphenophyte. | |
| Neocalamostachys | N. sharovii | Strobili of a large sphenophyte, presumably Neocalamites. | |
| "Pecopteris" | "P." filatovae | Marattiacean fern fronds | |
| Pleuromeiopsis | P. kryshtofovichii | Stems or branches of a lycopsid, up to in diameter. Nearly all fossils assigned to this species actually belong to other plants, but the holotype indicates that it is a legitimate and unique species similar to Ferganodendron but with larger leaf scars. | |
| Prynadaia | P. madygenica | A sphenophyte |
Pteridosperms ("seed ferns")
Pteridosperms are abundant and diverse, making up more than half of all the plant fossils recovered from the formation.| Genus | Species | Notes | Images |
| Edyndella | E. nikuzae | Leafy peltasperm shoots. | |
| Lepidopteris | L. ferganensis | Peltasperm fronds | |
| Lepidopteris | L. parvula | Peltasperm fronds | |
| Madygenia | M. asiatica | Peltasperm fronds, an endemic species. | |
| Madygenopteris | M. irregularis | Peltasperm fronds, an endemic species. | |
| Peltaspermum | P. madygenicum | Fertile shoots and braches of a peltasperm, bearing disc-shaped seed organs. | |
| Ptilozamites | P. davidovii | Enigmatic leaves of a possible pteridosperm | |
| Ptilozamites | P. elegans | Enigmatic leaves of a possible pteridosperm | |
| Sagenopteris | S. vakhrameevii | Caytonialean fronds | |
| Scytophyllum | S. pinnatum | Peltasperm fronds | |
| Uralophyllum | U. magnifolium | Peltasperm fronds | |
| Uralophyllum | U. petiolatum | Peltasperm fronds | |
| Uralophyllum | U. radczenkoi | Peltasperm fronds | |
| Uralophyllum | U. ramosum | Peltasperm fronds | |
| "Thinnfeldia" | "T." rhomboidalis | Frond-like leaves similar to some corystosperm foliage | |
| Vittaephyllum | V. bifurcatum | Peltasperm fronds | |
| Vittaephyllum | V. brickianum | Peltasperm fronds | |
| Vittaephyllum | V. ferganense | Peltasperm fronds | |
| Vittaephyllum | V. hirsutum | Peltasperm fronds |
Other gymnosperms
Though subordinate to pteridosperms in terms of abundance, other gymnosperms are still quite diverse, and some of them are far from uncommon.| Genus | Species | Notes | Images |
| Baiera | B. sp. | Ginkgoalean leaves | |
| Borysthenia | B. sp. | Conifer cones with associated branches and thin leaves. | |
| Ctenopteris | C. punctata | A dubious fern-like plant | |
| Cycadocarpidium | C. sp. | Seed-bearing cones of Podozamites-type conifers | |
| Ginkgoites | G. taeniatus | Ginkgoalean leaves | |
| Glossophyllum | G. ereminae | Enigmatic fronds of strap-shaped leaves, presumably related to Ginkgoales. | |
| Leuthardtia | L. sp. | Pollen-bearing cones, potentially from bennettitaleans | |
| Otozamites | O. sp. | Bennettitalean fronds | |
| Podozamites | P. distans | Conifer fronds with broad, strap-shaped leaves. | |
| Pseudoctenis | P. lanei | Cycad fronds | |
| Pterophyllum | P. hanesianum | Bennettitalean fronds | |
| Pterophyllum | P. firmifolium | Bennettitalean fronds | |
| Pterophyllum | P. pachartense | Bennettitalean fronds | |
| Pterophyllum | P. pinnatifidum | Bennettitalean fronds | |
| Rhaphidopteris? | R.? brickianae | Fronds of uncertain affinities, potentially among corystosperms, cycads, or ginkgoaleans. | |
| Sphenobaiera | S. granulifer | Ginkgoalean leaves | |
| Sphenobaiera | S. aff. zalesskyi | Ginkgoalean leaves | |
| Swedenborgia | S. cryptomerioides | Seed-bearning conifer cones | |
| Taeniopteris | T. latecostata | Gymnosperm leaves, possibly from cycads or bennettitaleans | |
| Taeniopteris | T. multinervia | Gymnosperm leaves, possibly from cycads or bennettitaleans | |
| Taeniopteris | T. spathulata | Gymnosperm leaves, possibly from cycads or bennettitaleans | |
| Taeniopteris | T.? stankevichii | Gymnosperm leaves, possibly from cycads or bennettitaleans | |
| Taeniopteridium | T. glossopteroides | Gymnosperm leaves, possibly from cycads or bennettitaleans | |
| Voltzia | V. sp. | Branches and needle-like leaves of voltzialean conifers |
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.