Vikinghøgda Formation Explained

Vikinghøgda Formation
Type:Formation
Age:Induan-Olenekian (Griesbachian-Spathian)
~
Period:Olenekian
Prilithology:Mudstone, siltstone
Otherlithology:Shale, sandstone
Region:Svalbard
Country:Norway
Paleocoordinates:42.2°N 10.4°W
Unitof:Sassendalen Group
Subunits:Deltadalen Member, Lusitaniadalen Member, Vendomdalen Member
Underlies:Botneheia Formation
Overlies:Kapp Starostin Formation

The Vikinghøgda Formation is a geologic formation in Svalbard, Norway. It preserves fossils dating back to the Early Triassic (Griesbachian-Spathian) period.[1] It is split into three members, from oldest to youngest: the Deltadalen Member (Induan), Lusitaniadalen Member (Smithian), and Vendomdalen Member (Spathian). The formation can be found in central Spitsbergen (Sassendalen, Dickson Land, James I Land), southern Spitsbergen, as well as the smaller islands of Barentsøya and Edgeøya. The type locality is positioned in the vicinity of Vikinghøgda and Sticky Keep, two low peaks along the southeast edge of Sassendalen (Sassen Valley) in Spitsbergen. The two upper members of the Vikinghøgda Formation were previously grouped together as the Sticky Keep Formation.[2] [3]

Subunits

The Vikinghøgda Formation is formed by fine-grained marine sediments, such as mudstones, shales, siltstones, and fine sandstones. There is a trend of finer sediments, deeper waters, and higher organic content through the formation. Though fossils are only abundant in the middle part (Lusitaniadalen Member) of the formation, index fossils can be found throughout the whole formation. The Vikinghøgda Formation is one of the better records of Early Triassic chronostratigraphy in the Boreal realm, owing to a combination of continuous sedimentation, distinctive index fossils, palynomorph stratigraphy, magnetostratigraphy,[4] and trace metal cyclostratigraphy.[5]

Deltadalen Member

The Deltadalen Member is named after the small stream valley which runs between Vikinghøgda and Sticky Keep. The member consists of about 70 meters of silty shales and fine sandstones, lying above the eroded surface of the Permian-age Kapp Starostin Formation. Some of the sandstone beds may be glauconitic or hummocky, and calcareous nodules may be present. Fossils are rare and restricted to these nodules, though they include a variety of silicified ammonoids, conodonts, bivalves, and gastropods.[6] Some index fossils have biostratigraphic significance, such as the ammonoid Otoceras boreale and the conodonts Neogondolella carinata and Neospathodus svalbardensis. These species constrain the member to the Induan stage, between the early Griesbachian and early Dienerian substages. The sandiest intervals can be found at the base and the top of the member. The depositional environment is reconstructed as a shallow marine setting influenced by storms and nearby deltaic sediments.

The Deltadalen Member has been equated with two other Induan-age formations in its vicinity: the Vardebukta Formation (exposed in western and southern Spitsbergen) and the Havert Formation (under the Barents Sea).

A 2020 study reported that the Permian-Triassic boundary occurs near the start of the Deltadalen Member. This was supported by several lines of evidence. The conodont Hindeodus parvus, which defines the base of the Triassic, was reported from a sediment layer 4.1 meters above the base of the member. An overlying tephra bed was dated to 252.13 ± 0.62 Ma via U-Pb radiometric dating, an age which is congruent with other reported estimates for the boundary.

Lusitaniadalen Member

The Lusitaniadalen Member is named after a glacier on the western flank of Vikinghøgda. The member is about 90 meters thick, mostly composed of laminated silty mudstones. Sandstone beds are less common, though when they occur, they are hummocky. Fossiliferous calcareous nodules are abundant in the sandstone-bearing layers, encasing well-preserved fossils of both invertebrates and vertebrates. The ammonoid index fossils Euflemingites romunderi and Wasatchites tardus indicate that the Lusitaniadalen Member was deposited during the Smithian substage of the Olenekian stage. The most common ammonoid is Arctoceras.[7] Fish and amphibian fossils have also been found in this member, particularly at the "Fish Niveau" bonebed. This horizon also produced the oldest known ichthyopterygian reptile fossil. The depositional environment corresponds to a deeper and calmer conditions, indicative of a major transgression (sea level rise) affecting the continental shelf.

The Lusitaniadalen Member has been equated with the Iskletten Member of the Tvillingodden Formation (in western Spitsbergen) and the lower part of the Klappmyss Formation (under the Barents Sea). It has also been previously described as the Iskletten Member of the Sticky Keep Formation.

Vendomdalen Member

Dark shale (laminated mudstone) dominates the 90-meter-thick Vendomdalen Member, though silty yellowish dolomitic beds also occur. Very large dolomite nodules or lenses, some over a meter in width, can also be found in this member. However, fossil-bearing concretions are rarer and more deformed than in the Lusitaniadalen Member. Fossils include abundant bivalves and Spathian ammonoid index fossils, such as Bajarunia euomphala, Keyserlingites subrobustus, and Parasibirites elegans. Marine reptile and fish bonebeds also occur in the Vendomdalen Member, the two most prominent being the older "Grippia Niveau" and younger "Lower Saurian Niveau".[8] [9] [10] [11] The Vendomdalen Member corresponds to the deepest part of the continental shelf, with high organic matter deposition and no influence from storm events.

The Vendomdalen Member has been equated with the Kaosfjellet Member of the Tvillingodden Formation (in western Spitsbergen) and the upper part of the Klappmyss Formation (under the Barents Sea). It has also been previously described as the Kaosfjellet Member of the Sticky Keep Formation.

Paleobiota

Tetrapods

Tetrapod remains occur in several horizons of the Vikinghøgda Formation. The fossils belong to ichthyosauriform and possibly sauropterygian reptiles and temnospondyl amphibians.

Reptiles

Reptiles of the Vikinghøgda Formation
Taxon / Genus Species Subunit Notes Images
CymbospondylusC. sp.Vendomdalen Member (Lower Saurian Niveau)A large basal ichthyosaur
Reptiliaindet.Vendomdalen Member (Grippia Niveau)An indeterminate reptile fossil (SVT 203) previously referred to Grippia longirostris. Postcranial proportions differ from all known ichthyosauromorphs, and may instead suggest affinities with Helveticosaurus or placodonts.[12]
GrippiaG. longirostrisVendomdalen Member (Grippia Niveau)A small basal ichthyopterygian
Ichthyopterygiaindet.Lusitaniadalen Member (Fish Niveau), Vendomdalen MemberIndeterminate ichthyopterygian remains. Pelagic-adapted ichthyopterygian fossils from the Lusitaniadalen Member are the oldest known from Svalbard and the world.[13]
IsfjordosaurusI. minorVendomdalen Member (Lower Saurian Niveau)An enigmatic ichthyosauromorph, possibly a hupehsuchian or basal ichthyopterygian
MerriamosaurusM. hulkeiVendomdalen Member (Lower Saurian Niveau)An ichthyopterygian based on forelimb fossils, initially described under the preoccupied name Rotundopteryx hulkei. Most likely a junior synonym of Pessopteryx nisseri.
OmphalosaurusO. nisseriVendomdalen Member (Lower Saurian Niveau)Weathered jaw fragments and other omphalosaurid-like fossils of uncertain validity, some of which have also been named as O. merriami or referred to Pessopteryx nisseri.
O. sp.Vendomdalen Member (Grippia and Lower Saurian Niveaus)Some fossils referred to Omphalosaurus appear to legitimately belong to the genus.[14]
PessopteryxP. nisseriVendomdalen Member (Lower Saurian Niveau)A large shastasaurid-like ichthyosaur, previously utilized as a chimera encompassing various ichthyopterygian fossils from the Lower Saurian Niveau.
QuasianosteosaurusQ. vikinghoegdaiVendomdalen Member (Grippia Niveau)A medium-sized ichthyopterygian
SvalbardosaurusS. crassidensVendomdalen Member (Grippia Niveau)A dubious tetrapod based on large teeth initially referred to ichthyosaurs, but more likely belonging to temnospondyl amphibians.

Amphibians

Temnospondyls of the Vikinghøgda Formation
Taxon / Genus Species Subunit Notes Images
AphanerammaA. rostratumLusitaniadalen Member (Fish Niveau)A long-snouted lonchorhynchine trematosaurid[15]
BoreosaurusB. thorslundiLusitaniadalen Member (Fish Niveau)A brachyopid or rhytidosteid
LyrocephaliscusL. euriLusitaniadalen Member (Fish Niveau)A trematosaurine trematosaurid, formerly known as Lyrocephalus.
PeltostegaP. ericiLusitaniadalen Member (Fish Niveau)A rhytidosteid
PlatystegaP. depressaLusitaniadalen Member (Fish Niveau)A trematosaurine trematosaurid
TertremaT. acutaLusitaniadalen Member (Fish Niveau)A relatively short-snouted trematosaurid, possibly a lonchorhynchine[16]
SassenisaurusS. spitzbergensisLusitaniadalen Member (Fish Niveau)Possibly an early capitosaur

Fish

A diverse fish assemblage has been described from the late Smithian[17] (early Olenekian) aged "Fish Niveau" (Lusitaniadalen Member, lower part of the "Sticky Keep Member") and other horizons, including lungfish, coelacanths, ray-finned fish, and hybodont sharks.[18] [10] [19] Most taxa were described by Erik Stensiö.[20] [21] The lists below are based on Kogan & Romano[17] and Bratvold et al.[10]

Lobe-finned fish

Sarcopterygii of the Vikinghøgda Formation
Taxon / Genus Species Subunit Notes Images
CeratodusC. sp.Vendomdalen Member (below Lower Saurian Niveau) Tooth plate of a lungfish
AxeliaA. robustaLusitaniadalen Member (Fish Niveau)A coelacanthid coelacanth
A. elegansLusitaniadalen Member (Fish Niveau)A coelacanthid coelacanth
MylacanthusM. lobatusLusitaniadalen Member (Fish Niveau)A coelacanthid coelacanth, quite similar to Axelia.
M. spinosusLusitaniadalen Member (Fish Niveau)A coelacanthid coelacanth, quite similar to Axelia.
SasseniaS. tuberculataLusitaniadalen Member (Fish Niveau)A rhabdodermatid coelacanth
S.? guttataLusitaniadalen Member (Fish Niveau)A rhabdodermatid coelacanth? Dubious generic attribution.[22]
S. sp.Lusitaniadalen Member (Fish Niveau)A rhabdodermatid coelacanth
ScleracanthusS. asperLusitaniadalen Member (Fish Niveau)A coelacanth, quite similar to Axelia.
WimaniaW. sinuosaLusitaniadalen Member (Fish Niveau)A relatively large coelacanthid coelacanth
W.? multistriataLusitaniadalen Member (Fish Niveau)A coelacanth
W.? sp.Lusitaniadalen Member (Fish Niveau)A coelacanth

Ray-finned fish

Actinopterygii of the Vikinghøgda Formation
Taxon / Genus Species Subunit Notes Images
AcrorhabdusA. bertiliLusitaniadalen Member (Fish Niveau)
A. asplundiLusitaniadalen Member (Fish Niveau)A ptycholepid
A. latistriatusLusitaniadalen Member (Fish Niveau)A ptycholepid
BirgeriaB. aldingeriLusitaniadalen Member (Fish Niveau)A birgeriid.The material was originally referred to B. mougeoti, the type species of Birgeria, but was later ascribed to its own species
B. sp.Lusitaniadalen Member (Fish Niveau)A partial skull of a large Birgeria[23]
BobasatraniaB. nathorstiLusitaniadalen Member (Fish Niveau)A large bobasatraniform that was originally referred to as Platysomus nathorsti
BoreosomusB. arcticusLusitaniadalen Member (Fish Niveau)A ptycholepid and the type species of Boreosomus, which was originally described as Acrolepis arctica by Arthur Smith Woodward
B. reuterskioldiLusitaniadalen Member (Fish Niveau)A ptycholepid
B.? scaberLusitaniadalen Member (Fish Niveau)
PteronisculusP. gyrolepidoidesLusitaniadalen Member (Fish Niveau)A poorly known rhadinichthyid or turseoid, It was originally described as Glaucolepis gyrolepidoides, but later referred to the genus Pteronisculus because the genus name Glaucolepis is preoccupied
PygopterusP. degeeriLusitaniadalen Member (Fish Niveau)A pygopterid. Pygopterus is one of the genera of ray-finned fish that crossed the Permian-Triassic boundary
SaurichthysS. elongatusLusitaniadalen Member (Fish Niveau) and possibly in beds slightly above itA saurichthyiform with a slender, elongated rostrum
S. hamiltoniLusitaniadalen Member (Fish Niveau)A saurichthyiform
S. ornatusLusitaniadalen Member (Fish Niveau)A saurichthyiform. This species is known primarily from skull remains
S. ornatus?Lusitaniadalen Member (Fish Niveau)A saurichthyiform. Based on one specimen that shows the postcranium and which has a different scale pattern than S. ornatus
S. wimaniLusitaniadalen Member (Fish Niveau)A saurichthyiform. Based on a specimen preserving the skull and anteriormost body portion that was originally described as "Belonorhynchus" wimani by Arthur Smith Woodward. A later found postcranium was referred to this species
S. sp.Lusitaniadalen Member (Fish Niveau)A saurichthyiform
TeffichthysT. woodwardiLusitaniadalen Member (Fish Niveau)A perleidiform. It was originally described as Colobodus altilepis by Arthur Smith Woodward but later referred to as Perleidus woodwardi (the species name was changed to avoid confusion with the type species of Perleidus, P. altolepis). The Early Triassic species of Perleidus were later ascribed to a different genus, Teffichthys[24]

Cartilaginous fish

Chondrichthyan remains are predominantely known from the Vendomdalen Member (Grippia Niveau). Several species listed below are also present in the nearby Vardebukta Formation, of Dienerian (late Induan) age.

Chondrichthyes of the Vikinghøgda Formation
Taxon / Genus Species Subunit Notes Images
AcrodusA. gaillardotiVendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
A. lateralisVendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
A. oppenheimeriVendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
A. scaberVendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
A. spitzbergensisVendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
A. vermiformisVendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
A. sp.Vendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
HybodusH. microdus/
H. ?microdus
Vendomdalen Member (Grippia Niveau)A hybodontid hybodontiform shark. Only known from isolated teeth
H. rapaxVendomdalen Member (Grippia Niveau)A hybodontid hybodontiform shark. Only known from isolated teeth
H. sasseniensisVendomdalen Member (Grippia Niveau)A hybodontid hybodontiform shark. Only known from isolated teeth
LissodusL. angulatusVendomdalen Member (Grippia Niveau)A lonchidiid hybodontiform shark. Only known from isolated teeth that were originally described as Polyacrodus angulatus
NemacanthusN. sp.Vendomdalen Member (Grippia Niveau)A palaeospinacid synechodontiform shark. Only known from fin spines
PalaeobatesP. polarisLusitaniadalen Member (Fish Niveau),
Vendomdalen Member (Grippia Niveau)
An acrodontid hybodontiform shark known from teeth and skelettal elements. Skull and postcranial elements are only known from the Fish Niveau[25]
P. sp.Vendomdalen Member (Grippia Niveau)An acrodontid hybodontiform shark. Only known from isolated teeth
PolyacrodusP. sp.Vendomdalen Member (Grippia Niveau)A polyacrodontid hybodontiform shark known from skull
HybodontoideaGen. et sp. indet.1Vendomdalen Member (Grippia Niveau)Basal plate of cephalic spine
Gen. et sp. indet.2Vendomdalen Member (Grippia Niveau)Fin spine
Gen. et sp. indet.3Vendomdalen Member (Grippia Niveau)Fin spine
SynechodontiformesGen. et sp. indet.1Vendomdalen Member (Grippia Niveau)Isolated teeth
Gen. et sp. indet.2Vendomdalen Member (Grippia Niveau)Isolated teeth

See also

Notes and References

  1. http://www.fossilworks.org/cgi-bin/bridge.pl?action=displayStrata&geological_group=&formation=Vikingh%F8gda&group_formation_member=Vikingh%F8gda Vikinghøgda Formation
  2. Mørk . Atle . Elvebakk . Geir . Forsberg . Arne W. . Hounslow. . Mark W. . Nakrem. . Hans Arne . Vigran . Jorunn Os . Weitschat . Wolfgang . 1999 . The type section of the Vikinghøgda Formation: a new Lower Triassic unit in central and eastern Svalbard . Polar Research . en . 18 . 1 . 51–82 . 10.1111/j.1751-8369.1999.tb00277.x . 2024-06-12 . 1999PolRe..18...51M . 0800-0395.
  3. Hurum . Jørn Harald . Engelschiøn . Victoria S. . Økland . Inghild . Bratvold . Janne . Ekeheien . Christina P. . Roberts . Aubrey J. . Delsett . Lene Liebe . Hansen . Bitten B. . Mørk . Atle . Nakrem . Hans Arne . Druckenmiller . Patrick S. . Hammer . Øyvind . 2018-10-04 . The history of exploration and stratigraphy of the Early to Middle Triassic vertebrate-bearing strata of Svalbard (Sassendalen Group, Spitsbergen) . Norwegian Journal of Geology . 98 . 2 . 165–174 . 10.17850/njg98-2-04. 216704366 . 10852/71101 . free .
  4. Hounslow . M. W. . Peters . C. . Mork . A. . Weitschat . W. . Vigran . J. O. . 2008-09-01 . Biomagnetostratigraphy of the Vikinghogda Formation, Svalbard (Arctic Norway), and the geomagnetic polarity timescale for the Lower Triassic . Geological Society of America Bulletin . en . 120 . 9–10 . 1305–1325 . 10.1130/B26103.1 . 2008GSAB..120.1305H . 0016-7606.
  5. Zuchuat . V. . Sleveland . A. R. N. . Twitchett . R. J. . Svensen . H. H. . Turner . H. . Augland . L. E. . Jones . M. T. . Hammer . Ø. . Hauksson . B. T. . Haflidason . H. . Midtkandal . I. . Planke . S. . 2020-09-15 . A new high-resolution stratigraphic and palaeoenvironmental record spanning the End-Permian Mass Extinction and its aftermath in central Spitsbergen, Svalbard . Palaeogeography, Palaeoclimatology, Palaeoecology . en . 554 . 109732 . 10.1016/j.palaeo.2020.109732 . 2020PPP...55409732Z . 218955779 . 0031-0182. 10852/81143 . free .
  6. Foster . William J. . Danise . Silvia . Twitchett . Richard J. . 2017-10-03 . A silicified Early Triassic marine assemblage from Svalbard . Journal of Systematic Palaeontology . en . 15 . 10 . 851–877 . 10.1080/14772019.2016.1245680 . 54030401 . 1477-2019. 2158/1152798 . free .
  7. Hansen . Bitten Bolvig . Bucher . Hugo . Schneebeli-Hermann . Elke . Hammer . Øyvind . The middle Smithian (Early Triassic) ammonoid Arctoceras blomstrandi: conch morphology and ornamentation in relation to stratigraphy . 2021 . Papers in Palaeontology . 7 . 3 . 1435–1457 . 10.1002/spp2.1348 . 2021PPal....7.1435H . free.
  8. Maxwell . Erin E. . Kear . Benjamin P. . 2013 . Triassic ichthyopterygian assemblages of the Svalbard archipelago: a reassessment of taxonomy and distribution . GFF . en . 135 . 1 . 85–94 . 10.1080/11035897.2012.759145 . 2013GFF...135...85M . 129092001 . 1103-5897.
  9. Hansen . Bitten Bolvig . Hammer . Øyvind . Nakrem . Hans Arne . 2018-10-04 . Stratigraphy and age of the Grippia niveau bonebed, Lower Triassic Vikinghøgda Formation, Spitsbergen . Norwegian Journal of Geology . 98 . 2 . 175–187 . 10.17850/njg98-2-02. 134204517 .
  10. Bratvold . Janne . Delsett . Lene Liebe . Hurum . Jørn Harald . 2018-10-04 . Chondrichthyans from the Grippia bonebed (Early Triassic) of Marmierfjellet, Spitsbergen . Norwegian Journal of Geology . 98 . 2 . 189–217 . 10.17850/njg98-2-03. 132293043 . 10852/71103 . free .
  11. Engelschiøn . Victoria Sjøholt . Delsett . Lene Liebe . Roberts . Aubrey Jane . Hurum . Jørn Harald . 2018-10-04 . Large-sized ichthyosaurs from the Lower Saurian niveau of the Vikinghøgda Formation (Early Triassic), Marmierfjellet, Spitsbergen . Norwegian Journal of Geology . 98 . 2 . 239–265 . 10.17850/njg98-2-05. 135275680 . 10852/71102 . free .
  12. Motani . Ryosuke . 2000 . Skull of Grippia longirostris : no contradiction with a diapsid affinity for the Ichthyopterygia . Palaeontology . en . 43 . 1 . 01–14 . 10.1111/1475-4983.00115. 2000Palgy..43....1M . 129452053 .
  13. Kear . Benjamin P. . Engelschiøn . Victoria S. . Hammer . Øyvind . Roberts . Aubrey J. . Hurum . Jørn H. . 2023 . Earliest Triassic ichthyosaur fossils push back oceanic reptile origins . Current Biology . en . 33 . 5 . R178–R179 . 10.1016/j.cub.2022.12.053. 36917937 . 257498390 . free .
  14. Ekeheien . Christina Pokriefke . Delsett . Lene Liebe . Roberts . Aubrey Jane . Hurum . Jørn Harald . 2018-10-04 . Preliminary report on ichthyopterygian elements from the Early Triassic (Spathian) of Spitsbergen . Norwegian Journal of Geology . 98 . 2 . 219–238 . 10.17850/njg98-2-07. 134421320 . 10852/71099 . free .
  15. Kear . Benjamin P. . Poropat . Stephen F. . Bazzi . Mohamad . 2016 . Late Triassic capitosaurian remains from Svalbard and the palaeobiogeographical context of Scandinavian Arctic temnospondyls . Geological Society, London, Special Publications . en . 434 . 1 . 113–126 . 10.1144/SP434.11 . 2016GSLSP.434..113K . 131531829 . 0305-8719.
  16. Slodownik . Miriam A. . Mörs . Thomas . Kear . Benjamin P. . 2021 . Reassessment of the Early Triassic trematosaurid temnospondyl Tertrema acuta from the Arctic island of Spitsbergen . Journal of Vertebrate Paleontology . en . 41 . 1 . e1900209 . 10.1080/02724634.2021.1900209 . 2021JVPal..41E0209S . 235574341 . 0272-4634.
  17. Kogan . Ilja . Romano . Carlo . 2016 . A new postcranium of Saurichthys from the Early Triassic of Spitsbergen . Freiberger Forschungshefte C (Paläontologie, Stratigraphie, Fazies 23) . 550 . 205–221 . 9783860125526.
  18. Cox . C. B. . Smith . David G. . 1973 . A review of the Triassic vertebrate faunas of Svalbard . Geological Magazine . en . 110 . 5 . 405–418 . 10.1017/S0016756800036190 . 1973GeoM..110..405C . 128809142 . 0016-7568.
  19. Brinkmann . Winand . Romano . Carlo . Bucher . Hugo . Ware . David . Jenks . Jim . 2010 . Palaeobiogeography and stratigraphy of advanced gnathostomian fishes (Chondrichthyes and Osteichthyes) in the Early Triassic and from selected Anisian localities (Report 1863–2009) . Zentralblatt für Geologie und Paläontologie, Teil II . 2009 . 765–812 . .
  20. Book: Stensiö, E. . 1921 . Triassic fishes from Spitzbergen 1 . xxviii+307 . Adolf Holzhausen . Wien.
  21. Stensiö . E. . 1925 . Triassic fishes from Spitzbergen 2 . Kungliga Svenska Vetenskapsakademiens Handlingar . 3 . 1–261.
  22. Book: Forey, Peter L. . 1998 . History of the coelacanth fishes . Chapman & Hall . London . 434 . 9780412784804.
  23. Scheyer . Torsten M. . Romano . Carlo . Jenks . Jim . Bucher . Hugo . 2014 . Early Triassic Marine Biotic Recovery: The Predators' Perspective . PLOS ONE . 9 . 3 . e88987 . 10.1371/journal.pone.0088987 . 24647136 . 3960099 . 2014PLoSO...988987S . free.
  24. Marramà, G. . Lombardo, C. . Tintori, A. . Carnevale, G. . amp . 2017 . Redescription of "Perleidus"(Osteichthyes, Actinopterygii) from the Early Triassic of northwestern Madagascar. . Rivista Italiana di Paleontologia e Stratigrafia . 123 . 2 . 219–242 . 10.13130/2039-4942/8328. free.
  25. Romano . Carlo . Brinkmann . Winand . A new specimen of the hybodont shark Palaeobates polaris with threedimensionally preserved Meckel's cartilage from the Smithian (Early Triassic) of Spitsbergen . Journal of Vertebrate Paleontology . December 2010 . 30 . 6 . 1673–1683 . 10.1080/02724634.2010.521962. 2010JVPal..30.1673R . 86411191 .