Madygen Formation
Madygen Formation | ||
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Ma | ||
Approximate paleocoordinates 41°12′N 60°36′E / 41.2°N 60.6°E | | |
Region | Batken & Osh Regions | |
Country | Kyrgyzstan Tajikistan Uzbekistan | |
Extent | Fergana Valley & Range | |
Type section | ||
Named for | Madygen village | |
Named by | Evgeny A. Kochnev | |
The Madygen Formation (
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
Description
![](http://upload.wikimedia.org/wikipedia/commons/thumb/3/3b/230_Ma_plate_tectonic_reconstruction.png/250px-230_Ma_plate_tectonic_reconstruction.png)
The Madygen Formation is a 560 metres (1,840 ft)[5] 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.[6] 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.[7][4]
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.[7][1] The fluvio-lacustrine deposits of the Madygen Formation belong to one of only a few occurrences of continental Triassic beds in Central Asia.[8]
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.[2] Though fossils can be found throughout the entire formation, the most extensive fossil deposits occur in the northern area, known as Dzaylyaucho (also spelled Dzhayloucho or Dzhailoucho), 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.[9][4][1]
The environment represented by the Madygen Formation was positioned on the
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.[6]
The hydrocarbon potential of samples of the Madygen Formation ranges from poor to excellent. The sediments containing more than 0.5%
Paleontological significance
During the 1960s, Russian paleontologists recovered an unusually rich fossil content in the type strata of the Madygen Formation, including abundant
Fish remains mostly represent endemic genera assigned to the actinopterygian families
Paleobiota
Color key
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Notes Uncertain or tentative taxa are in small text; |
Amphibians
Genus | Species | Material | Notes | Images |
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Madygenerpeton[12] | M. pustulatus[12] | Skull and osteoderms. | A reptiliomorph with aquatic adaptations. It was first named in 2010 on the basis of a nearly complete skull and associated osteoderms. It seems that this tetrapod could've been semi-aquatic, due to the increased flexibility of the trunk region.[12][13]
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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).[14] | ![]() |
Reptiles
Reptiles are the most abundant of the tetrapod paleofauna at Madygen, with three described genera.
Genus | Species | Material | Notes | Images |
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Kyrgyzsaurus[15] | K. bukhanchenkoi[15] | A single specimen preserving the front half of a skeleton and scale impressions.[16][15] | Potentially the oldest known member of the drepanosaurs, a diverse clade of Triassic reptiles that evolved arboreal and fossorial lifestyles.[17] It was also the first member of the group to be described from Asia.[15] | ![]() |
Longisquama[18] | L. insignis[18] | A specimen preserving the front half of a skeleton and "plumes", and at least five additional "plume" fragments.[3] | A neodiapsid reptile possessing unusual large scaly growths ("plumes") 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.[18][3][19]
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Sharovipteryx | S. mirabilis[20] | A single skeleton with impressions of gliding membranes, split across a slab and counterslab.[3] | A gliding archosauromorph, and the type genus of the family Sharovipterygidae. Originally named as Podopteryx,[20] a genus name which was preoccupied by a damselfly.[21][3] | ![]() |
Synapsids
Genus | Species | Material | Notes | Images |
---|---|---|---|---|
Madysaurus[22] | M. sharovi[22] | A single partial skeleton. | A small | ![]() |
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.[23][2]
Genus | Species | Material | Notes | Images |
---|---|---|---|---|
Fayolia | F. sharovi[2] | Egg capsules | An egg capsule likely belonging to a | |
Palaeoxyris | P. alterna[2] | Egg capsules | The egg cases of hybodonts that appear frequently throughout the fossil record.[2] 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[2] | Teeth, denticles,[23] and egg capsules | A hybodontid elasmobranch 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. 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.[2]
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Bony fishes
The following fish fossils were found in the formation:[25][9][26][23][1]
Genus | Species | Material | Notes | Images |
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Alvinia[25] | A. serrata[25] | Partial skeletons. | A small | |
Asiaceratodus | A. sharovi | Partial skeletons. | A medium-sized dipnoan (lungfish), around 30 centimetres (12 in) in length.[9][26] | |
Ferganiscus[25] | F. osteolepis[25] | Nearly complete skeletons. | A small and abundant palaeoniscid, 10–20 centimetres (3.9–7.9 in) in length.[9][26][1] | |
Megaperleidus[25] | M. lissolepis[25] | Partial skeletons. | A medium-sized perleidid.[9][26] | |
Oshia[25] | O. ferganica[25] | Partial skeletons, scales.[23] | A medium-sized scanilepiform with predatory habits. Around 45 centimetres (18 in) in length.[26]
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Saurichthys | S. orientalis[25] | Nearly complete skeleton,[26] scales,[23] other fragmentary material. | A relatively small saurichthyid chondrostean, around 45 centimetres (18 in) in length.[26]
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Sixtelia[25] | S. asiatica[25] | Nearly complete skeletons, scales.[23] | A small and common palaeoniscid, 6–12 centimetres (2.4–4.7 in) in length.[9][26][1] |
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.[27] Under some estimates, it has the most diverse insect assemblage of the entire Permian-Triassic interval.[28][4] From EDNA (The Fossil Insect Database), Shcherbakov (2008),[4] and the Paleobiology Database, unless cited otherwise:
Odonata (damselflies, dragonflies and kin)
The Odonata and other Odonatoptera of the Madyen Formation are rare (about 100 specimens), with moderate diversity.[4] They were described most extensively by Pritykina (1981).[29]
- †Paurophlebiidae (8 species): Cladophlebia (C. brevis, C. parvula), Neritophlebia (N. elegans, N. longa, N. vicina), Nonymophlebia (N. venosa), Paurophlebia (P. angusta, P. lepida)
- †Triadophlebiidae (6 species): Triadophlebia (T. distincta, T. honesta, T. madygenica, T. magna, T. minuta, T. modica)
- †Zygophlebiidae (4 species): Cyrtophlebia (C. sinuosa), Mixophlebia (M. mixta), Zygophlebia (Z. ramosa), Zygophlebiella (Z. curta)
- †Kennedyidae (4 species): Kennedya (K. carpenteri, K. ferganensis,[30] K. gracilis, K. madygensis[30])
- †Voltzialestidae (3 species):[31] Terskeja (T. paula, T. pumilio, T. tenuis)
- †Batkeniidae (2 species): Batkenia (B. pusilla), Paratriassoneura[32] (P. primitiva)
- †Mitophlebiidae (1 species): Mitophlebia (M. enormis)
- †Protomyrmeleontidae (1 species): Ferganagrion (F. kirghiziensis[33])
- †Triadotypidae (1 species): Reisia (R. sogdiana[32])
- †Triassolestidae (1 species): Triassolestodes (T. asiaticus)
- †Xamenophlebiidae (1 species): Xamenophlebia (X. ornata)
Blattodea (roaches)
The Blattodea of the Madygen Formation are extremely abundant (a quarter of all insect fossils collected), 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.[34][4] Some species were given a preliminary description by Vishniakova (1998).
- †Caloblattinidae: Sogdoblatta (S. maxima, S. nana, S. porrecta),[35] Thuringoblatta (T. sogdianensis)
- †Subioblattidae: Subioblatta (S. madygenica[36])[27]
- Undescribed species of †Blattulidae, †Phyloblattidae, †Poroblattinidae, †Spiloblattinidae, and potentially †Archimylacridae.
†Titanoptera
![](http://upload.wikimedia.org/wikipedia/commons/thumb/a/a2/Gigatitan_vulgaris.jpg/141px-Gigatitan_vulgaris.jpg)
The †Titanoptera of the Madygen Formation are uncommon (about 200 specimens), with moderate diversity, though still higher than any other geological formation.[4] They were described most extensively by Sharov (1968) and Gorochov (2003). 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 (16 in).[37] 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.[38]
- †Paratitanidae (11 species): Microtitan (M. zherichini), Paratitan (P. bispeculum, P. intermedius, P. latispeculum, P. libelluloides, P. longispeculum, P. modestus, P. ovalis, P. reductus, P. reliquia,[39][40] P. venosus)
- †Mesotitanidae / †Gigatitanidae (11 species): 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 suborder of orthopterans, rather than its own order. Under this system, the diversity of Madygen titanopterans is also reduced to seven species spread out across two genera (Mesotitan and Gigatitan).[41]
Orthoptera (grasshoppers and kin)
The Orthoptera of the Madygen Formation are very common (about 1400 specimens) and very diverse (over 100 species). Most species belong to the suborder Ensifera (kin of crickets, katydids, and grigs), with only a few Caelifera (kin of grasshoppers). None of the families survive to the present, and most are endemic to the Triassic.[4] Madygen orthopterans were described most extensively by Sharov (1968) and Gorochov (1986, 1987, 1994, 2005).
- †Haglidae (44 species): 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),[42] 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 species): Batkenella (B. megaptera), Brevibatkenella (B. abscisa), Dolichobatkenella (D. perlonga), Eubatkenella (E. devexia), Gryllacrimima (G. elongata, G. madygenioides, G. perfecta, G. simplicis), Kashgarlimahmutia (K. reducta),[43][42] Madygenia (M. extremalis, M. longissima, M. orientalis, M. ovalis), Oedischimima (O. deficientis), Parafergania (P. sharovi), Platymadygenia (P. grandis)
- †Xenopteridae (15 species): 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 species): Brevilocustavus (B. distinctus, B. microscopicus), Ferganopsis (F. lanceolatus), Locustavus (L. deformatus, L. intermedius, L. madygensis, L. minutus, L. problematicus), Miolocustavus (M. reductus)
- †Bintoniellidae (8 species): 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 species): Adzhajloutshella (A. planis, A. talis), Dzhajloutshella (D. arcanum, D. flexuosa), Triassoxya (T. novozhilovi)
- †Tuphellidae (4 species): Neotuphella (N. minor), Tuphella (T. rasnitsyni, T. rohdendorfi, T. sharovi)
- †Gryllavidae (3 species): Gryllavus (G. madygenicus), Paragryllavus (P. curvatus), Zaragryllavus (Z. elongatus)
- †Mesoedischiidae (3 species): Mesoedischia (M. kirgizia, M. madygenica, M. obliqua)
- †Permelcanidae (2 species): Meselcana (M. madygenica, M. permelcanoides)
- †Hagloedischiidae (1 species): Hagloedischia (H. primitiva)
Phasmatodea (stick insects and kin)
The Phasmatodea of the Madygen Formation are somewhat common (about 350 specimens) and somewhat low in diversity. All of the families were endemic to the Triassic.[4] They were described most extensively by Sharov (1968) and Gorochov (1994).
- †Prochresmodidae (8 species): Prochresmoda (P. longipoda, P. media, P. minuta, P. parva), Triassophasma (T. brevipoda, T. intermedium, T. minutissimum, T. pusillum)
- †Aeroplanidae (2 species): Sharovoplana (S. affinis, S. parallelica)[42]
- †Xiphopteridae? (2 species): Xiphopterum (X. curvatum, X. sharovi)
Grylloblattodea (ice-crawlers and kin) and other basal polyneopterans
The
- †Megakhosaridae (11 species):[44] Madygenocephalus (M. micropteron[45]), Megablattogryllus (M. austerus, M. magister, M. pinguis), Megakhosarodes (M. obtusus, M. paulovenosus), Mesoblattogryllus (M. abruptus,[45] M. intermedius), Metakhosara (M. sharovi), Protoblattogryllus (P. asiaticus, P. variabilis)
- †Mesorthopteridae (11 species): Belmophenopterum (B. rasnitsyni[46]), Austroidelia (A. asiatica), Locustoblattina (L. marginata,[47] L. segmentata[47]), Mesoidelia (M. faceta, M. rasnitsyni,[48] M. semota), Parastenaropodites (P. fluxa,[47] P. longiuscula, P. nervosa), Sharovites (S. alexanderi[49])
- †Blattogryllidae (9 species):[44] Anoblattogryllus (A. fundatus), Baharellinus (B. dimidiatus, B. pectinatus), Baharellus (B. madygensis), Costatoviblatta (C. aenigmatosa, C. conjuncta,[44] C. longipennis,[44] C. similis[45]), Dorniella (D. primitiva)
- †Gorochoviidae (8 species): Gorochovia (G. anomala, G. bifurca, G. fecunda, G. individua, G. minuta), Gorochoviella (G. conjuncta), Pseudoliomopterites (P. lucidus, P. obscurus)
- †Ideliidae (7 species): Anaidelia (A. extrema), Ideliopsina (I. nana, I. ornata, I. ruginosa, I. stupenda), Madygenidelia (M. conjuncta), Pseudoshurabia (P. pallidula)
- †Geinitziidae (5 species): Geinitziella (G. rasnitsyni[50]), Shurabia (S. anomala, S. ferganensis, S. minutissima,[50] S. tanga[51])
- †Madygenophlebiidae (4 species): Madygenophlebia (M. bella, M. nana, M. primitiva), Micromadygenophlebia (M. obscura)
- †Sylvabestiidae (2 species): Aiban (A. kichineis[52]), Sharovala (S. triassica[53])
- †Necrophasmatidae (1 species):[54] Ferganamadygenia (F. plicata)
- †Daldubidae (1 species): Batkentak (B. intactus[55])
- †Sylvaphlebiidae (1 species): Batkenopterum (B. kirgizicum[53])
†Miomoptera
The †Miomoptera of the Madygen Formation are somewhat common (about 500 specimens),[4] most or all of which belong to a single valid species.[56][57]
- †Permosialidae (1 species): Permosialis (P. triassica[58])
Plecoptera (stoneflies)
The Plecoptera of the Madygen Formation are rare (about 80 specimens) and somewhat low in diversity.[4] They were described most extensively by Sinitchenkova (1987).
- †Perlariopseidae (14 species): 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[59]), Tritaniella (T. mera, T. pectinata, T. perlonga, T. synneura)
- †Siberioperlidae (1 species): Siberioperla (S. ovalis)
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.[47]
- †Alexarasniidae: Nestorembesia (N. novojilovi,[60] N. shcherbakovi[47])
- †Rasnalexiidae: Rasnalexia (R. rasnitsyni[61])
Hemiptera (true bugs)
The
- †Maguvopseidae:[62] Asiocula (A. lima[62]), Cuanoma (C. protracta[62]), Falcarta (F. bella[62]), Fasolinka (F. beckermigdisovae[62]), Krendelia (K. ansata[62]), Maguviopsis (M. kotchnevi), Nevicia (N. imitans[62]), Nonescyta (N. mala[62]), Phyllotexta (P. latens[62]), Sacvoyagea (S. ventrosa[62]), Sitechka (S. perforata[62])
- †Palaeontinidae: Papiliontina (P. dracomima,[63] P. machaon,[63] P. spectans[63])
- †Mesojabloniidae: Fulgobole (F. evansi[62]), Mesojablonia (M. kukalovae), Scytachile (S. emeljanovi[62])
- †Dunstaniidae: Dunstaniodes (D. elongatus), Siksteliana (S. popovi)
- †Curvicubitidae: Beaconiella (B. cincta,[64] B. pulchra[64])
- †Creaphididae: Creaphis (C. theodora)
- †Ipsviciidae: Strivicia (S. davidi[65])
- †Naibiidae: Coccavus (C. supercubitus[66])
- †Paraknightiidae: Triknightia (T. mira[64])
- †Progonocimicidae: Pelorisca (P. connectens)
- †Saaloscytinidae:[62] Tingiopsis (T. reticulata)
- †Serpentivenidae: Serpentivena (S.tigrina)
- Undescribed species of †Ochteroidea.
Neuroptera (lacewings and kin)
The Neuroptera of the Madygen Formation are uncommon (around 200 specimens) but diverse in form. Only a few species have been formally described.[4][67]
- †Archeosmylidae: Madygoneura (M. elongata[67]), Osmylotriasia (O. superba[67]), Triasella (T. ovata[67])
- Berothidae (beaded lacewings): Ferganoberotha (F. miniutissima[68])
- †Permithonidae: Relictovia (R. pristina[67])
- Undescribed species of †Polystoechotidae" (giant lacewings).
Coleoptera (beetles)
The
- †Schizophoridae (17 species): 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 species):[69] 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 species):[69] Lithocupes (L. gigas, L. incertus, L. punctatus), Notocupes?[70] (N. laticella, N. rostratus, N. tenuis), Notocupoides? (N. capitatus, N. fasciatus, N. triassicus), Rhabdocupes? (R. baculatus, R. longus, R. minor)
- †Ademosynidae (7 species):[71] Ademosyne (A. bacca, A. elliptica, A. kirghizica), Cephalosyne (C. capitata), Dolichosyne (D. confragosa, D. rostrata, D. sulcata)
- †Triaplidae (7 species):[72] Avocatinus (A. elongatus), Catinoides (C. rotundatus), Macrocatinius (M. brachycephalus), Triaplus (T. laticoxa, T. macroplatus), Triassocatinius (T. brachynotus, T. glabratus)
- †Obrieniidae (5 species): Guillermia (G. lecticula), Madygenorhynchus (M. multifidus), Obrienia (O. illaetabilis, O. ingurgata, O. kuscheli)
- †Peltosynidae (4 species):[30] Gnathopeltos (G. dixis[30]), Ofthalmopeltos (O. synkritos[30]), Peltosyne (P. triassica, P. varyvrosa[30])
- †Asiocoleidae[73] / †Tricoleidae (3 species): Sogdelytron (S. latum), Tricoleodus (T. acutus, T. longus)
- sensu stricto / Cupedinae (reticulated beetles, 2 species): Mesocupoides(M. indistinctus, M. proporeius)
- Trachypachidae (false ground beetles, 1 species): Sogdodromus (S. altus)
Hymenoptera (wasps and kin)
The Hymenoptera of the Madygen Formation are rare (about 60 specimens), but with moderate diversity.[4] All the species belong to the sawfly family Xyelidae, which still survives to the present. They were described most extensively by Rasnitsyn (1964, 1969).[74]
- Xyelidae (xyelid sawflies, 36 species): Asioxyela (A. parvula, A. paurura, A. smilodon), Chubakka (C. madygensis[75]), 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,[75] M. bashkuevi,[75] M. kurochkini,[75] M. levivenosa[75]), Madygenius (M. extraradius, M. primitivus), Oryctoxyela (O. anomala, O. triassica), Microxyela (M. minuta[76]), Samarkandykia (S. ryzhkovae,[75] S. shmakovi[75]), Triassoxyela (T. foviolata, T. grandipennis, T. orycta, T. sharovi[76]), Xiphoxyela (X. procrusta, X. striata), Xyelinus (X. angustiradius, X. major, X. scherbachov[76])
Mecoptera (scorpionflies)
The Mecoptera of the Madygen Formation are very common (about 1600 specimens) and fairly diverse (over 40 species).[4] They were described most extensively by Ponomarenko & Rasnitsyn (1974), and Novokshonov (1997, 2001).
- †Permochoristidae (15 species):[34] 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 species):[34] 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 species): Blattomerope (B. polyneura), Pronotiothauma (P. neuropteroides), Thaumatomerope (T. madygenica, T. minuta, T. oligoneura, T. sogdiana)
- †Mesopsychidae (5 species): Mesopsyche (M. gentica, M. justa, M. ordinata, M. shcherbakovi, M. tortiva)
- incertae sedis (2 species): †Mecaenigma (M. suspectum, M. viduum)
- †Pseudopolycentropodidae (1 species): Pseudopolycentropus (P. madygenicus)
Trichoptera (caddisflies)
The
- †Cladochoristidae: Cladochorista (C. curta[77]), Cladochoristella (C. sola[77])
- †Prorhyacophilidae: Prorhyacophila (P. batkenica,[77] P. furcata, P. rara,[77])
- †Necrotauliidae: Paranecrotaulius (P. proximus)[78]
- Philopotamidae? Prophilopotamus (P. asiaticus) (dubious)[34]
Diptera (flies)
The
- †Psychotipidae (2 species):[80] Psychotipa (P. depicta, P. predicta)
- †Protorhyphidae (2 species): Vymrhyphus (V. triassicus, V. tuomikoskii)
- †Gnomuscidae? (2 species):[80] Gnomusca (G. molecula, G. renyxa)
- Chaoboridae (phantom midges, 1 species): Triassomyia (T. shcherbakovi[81])
- †Hennigmatidae? (1 species): Anemeca (A. liya)
- †Kuperwoodidae (1 species):[80] Kuperwoodia (K. benefica)
- Limoniidae (limoniid crane flies, 1 species): Mabelysia (M. charlesi)
- †Nadipteridae (1 species): Nadiptera (N. pulchella)
- †Vladipteridae (1 species): Dilemmala (D. specula)
Other insects
Other invertebrates
Apart from insects, the most common invertebrates found in the Madygen Formation are
Flora
The plant fossils of the Madygen Formation were reviewed in detail by Dobruskina (1995).[7] 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.[84]
Non-vascular plants
Genus | Species | Notes | Images |
---|---|---|---|
Muscites | M. brickiae | A moss-like bryophyte with long, slender leaves akin to some species of Cryptopodium. Another unnamed species of Muscites is also found in the Madygen Formation.[85] | |
Ricciopsis | R. ferganica | A | |
Thallites | T. sp. | Large leaf-like "thallophytes" (algae or other simply plant-like organisms) of uncertain affinities, some up to 25 centimetres (9.8 in) long.[7] |
Ferns, horsetails, and lycopods
Genus | Species | Notes | Images |
---|---|---|---|
Chiropteris | C. integalla | Fern fronds | |
Cladophlebis | C. ex. gr. nebbensis | Osmundacean fern fronds | |
C. ex. gr. paralobifolia | Osmundacean fern fronds | ||
C. raciborskii | Osmundacean fern fronds | ||
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 (horsetail)
|
|
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 30 centimetres (12 in) in diameter,[7] but most fossils suggest a much smaller maximum diameter of 5.5 centimetres (2.2 in).[86] Previously misidentified as a species of Sigillaria. | |
Isoetites | I. madygensis | Quillwort corms (bulb-like stems) with attached leaves, very similar to modern Isoetes. Also probably includes lycopod scales previously named as Annalepis leae.[86] 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.[87]
|
|
I. sixteliae | Serrated quillwort leaves, quite common.[86] | ||
Mesenteriophyllum | M. kotschnevii | Enigmatic strap-like leaves, up to 14 centimetres (5.5 in) 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 (most likely a misinterpretation of insect damage), 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 (Fraser in 2006). However, Buchwitz & Voigt (2012) 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.[86][19] | |
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 6 centimetres (2.4 in) 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.[86] | |
Prynadaia | P. madygenica | A sphenophyte |
Pteridosperms ("seed ferns")
Genus | Species | Notes | Images |
---|---|---|---|
Edyndella | E. nikuzae | Leafy peltasperm shoots. | |
Lepidopteris | L. ferganensis | Peltasperm fronds | |
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 | |
P. elegans | Enigmatic leaves of a possible pteridosperm | ||
Sagenopteris | S. vakhrameevii | Caytonialean fronds | |
Scytophyllum | S. pinnatum | Peltasperm fronds | |
Uralophyllum | U. magnifolium | Peltasperm fronds | |
U. petiolatum | Peltasperm fronds | ||
U. radczenkoi | Peltasperm fronds | ||
U. ramosum | Peltasperm fronds | ||
"Thinnfeldia" | "T." rhomboidalis | Frond-like leaves similar to some corystosperm foliage | |
Vittaephyllum | V. bifurcatum | Peltasperm fronds | |
V. brickianum | Peltasperm fronds | ||
V. ferganense | Peltasperm fronds | ||
V. hirsutum | Peltasperm fronds |
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.[7][88]
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[10] | |
Podozamites | P. distans | Conifer fronds with broad, strap-shaped leaves. | |
Pseudoctenis | P. lanei | Cycad fronds[10] | |
Pterophyllum | P. hanesianum | Bennettitalean fronds | |
P. firmifolium | Bennettitalean fronds[10] | ||
P. pachartense | Bennettitalean fronds | ||
P. pinnatifidum | Bennettitalean fronds[10] | ||
Rhaphidopteris? | R.? brickianae | Fronds of uncertain affinities, potentially among corystosperms, cycads, or ginkgoaleans. | |
Sphenobaiera | S. granulifer | Ginkgoalean leaves | |
S. aff. zalesskyi | Ginkgoalean leaves | ||
Swedenborgia | S. cryptomerioides | Seed-bearning conifer cones | |
Taeniopteris | T. latecostata | Gymnosperm leaves, possibly from cycads or bennettitaleans | |
T. multinervia | Gymnosperm leaves, possibly from cycads or bennettitaleans | ||
T. spathulata | Gymnosperm leaves, possibly from cycads or bennettitaleans | ||
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.[11] 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.[11]
See also
- Carnian formations
- Santa Maria Formation, highly fossiliferous formation in Rio Grande do Sul, Brazil
- Denmark Hill Insect Bed, insect-bearing unit of Queensland, Australia
- Candelária Formation, lacustrine formation of the Paraná Basin, Brazil
- Ischigualasto Formation, alluvial and fluvial Lagerstätte of Argentina
- Los Rastros Formation, insect-bearing formation underlying the Ischigualasto Formation
- Doswell Formation, continental fossiliferous formation of the eastern United States
- Lossiemouth Sandstone, fossiliferous formation of Scotland
- Other Central Asian Lagerstätten
- Bissekty Formation, Turonian formation of Uzbekistan
- Bostobe Formation, Santonian to Campanian formation of Kazakhstan
- Karabastau Formation, Late Jurassic formation of Kazakhstan
- Danata Formation, Paleogene formation of Turkmenistan
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Further reading
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. .