Mochlodon
| Mochlodon | ||||||||||||
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Mochlodon suessi ; Lectotype PIUW 2349/2 |
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| Temporal occurrence | ||||||||||||
| Upper Cretaceous ( Santonium - Lower Campanium ) | ||||||||||||
| 86.3 to 76 million years | ||||||||||||
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| Scientific name | ||||||||||||
| Mochlodon | ||||||||||||
| Seeley , 1881 | ||||||||||||
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Mochlodon is a genus of small- sized pelvic dinosaurs from the group of the Rhabdodontidae within the Iguanodontia . The genus is known with two species from the Upper Cretaceous of Austria and Hungary .
Etymology and history of research
The generic name Mochlodon is derived from the ancient Greek terms Μοχλός (" mokhlos ": "lever", "rod", "pole") and ὀδών (" odon ": "tooth").
In 1859 Eduard Suess and his then pupil Ferdinand Stoliczka searched the dump material of the coal mine near Felbring ( Muthmannsdorf ) in Lower Austria for Cretaceous freshwater mollusks . Stoliczka succeeded in finding a small tooth, the shape of which was strongly reminiscent of that of the Iguanodon dinosaur genus described by Gideon Mantell in 1824 .
Attempts by the mountain administrator Pavlovich to narrow down the horizon more precisely and to retrieve further vertebrate fossils were initially unsuccessful. Only when driving the new "Constantine tunnel" the fossil-bearing layer was ascended again and you could collect substantial evidence that at the Paleontological Institute of the University of Vienna was passed. Emanuel Bunzel was commissioned with an initial inspection and description of the finds. His efforts were published in 1871 and resulted, among other things, in the first description of the genus Struthiosaurus . Bunzel described the " Iguanodon -like" finds from this collection on the basis of a lower jaw fragment (PIUW 2349/2), a single tooth and two vertebral bodies , in honor of Suess, initially as Iguanodon suessii .
In the spring of 1879, Suess invited the British paleontologist Harry Govier Seeley to come to Vienna and examine the Muthmannsdorf finds more closely. Seeley spent about a month in Vienna and came to significantly different conclusions than Bunzel , especially with regard to Iguanodon suessii . On the one hand, he saw sufficient differences to the genus Iguanodon in the lower jaw fragment (PIUW 2349/2) to justify the establishment of a new genus Mochlodon ; on the other hand, he interpreted other finds that Bunzel had assigned to various other reptile groups as probably belonging to Mochlodon suessii . Remarkably, in his new description published in 1881, Seeley only made comparisons with the genus Iguanodon , but not with the genus Rhabdodon from the French Cretaceous described by Philippe Matheron in 1869 .
At the beginning of the 20th century, Franz von Nopcsa described similar finds from the area around Hațeg in today's Romania in a series of publications , which he initially referred to as Mochlodon suessi or Mochlodon (?) Robustum . In 1902 Nopcsa revised his opinion and interpreted Mochlodon suessi as juveniles of Mochlodon robustum , whereby the latter taxon became obsolete as a junior synonym of Mochlodon suessi . After a direct comparison with the evidence from Rhabdodon priscus , which Matheron had already described in 1869 but not accurately depicted, Nopcsa came to the conclusion in 1915 that Rhabdodon priscus and Mochlodon suessi were identical and that the morphological differences were due to a sexual dimorphism . Consequently, Nopcsa only differentiated in this work between Rhabdodon priscus and Rhabdodon priscus var. Suessi , which he interpreted as different sexes of the same species. In 1923 Nopcsa tried to substantiate his hypothesis, synonymized Mochlodon with Rhabdodon priscum and assigned the French evidence and the Romanian forms originally called Mochlodon robustum to a female gender, while he interpreted the forms previously described as Mochlodon suessi as male individuals of the same species. In 1929, however, he claimed the exact opposite and designated Rhabdodon priscum (= Rhabdodon robustum ) as a male and Rhabdodon suessi as a female of the same species.
After Nopcsa's death in 1933, basic research on this topic came to a virtual standstill. However, the importance of the finds was generally recognized and was included in numerous specialist publications and textbooks. However, many of these authors continued to use the generic name Mochlodon , which was because the name Rhabdodon was strictly invalid in this context. As early as 1831 Friedrich Ludwig Fleischmann had described an allegedly new species of snake as Rhabdodon fuscus . Although the snake described and depicted by Fleischmann could later easily be identified as a representative of the lizard snakes ( Malpolon ), a taxon that Leopold Fitzinger had already introduced in 1826, the name Rhabdodon was clearly documented and already in 1869, when Matheron described the French fossil finds not available.
In order to solve the problem, Winand Brinkmann applied to the ICZN in 1986 for the taxon Rhabdodon Fleischmann to be deleted from the official list of zoological generic names in 1831 and instead to include the taxon Rhabdodon Matheron , 1869 in the list ("Case 2536"). The motion came to the vote at the ICZN in 1987 and was approved by a large majority ("Opinion 1483"). The result of the vote was published in 1988 and thus all relevant finds from France, Austria and Romania were assigned to the genus Rhabdodon .
In 2003 David B. Weishampel and co-authors separated the Romanian finds as an independent genus Zalmoxes with two species ( Zalmoxes robustus and Zalmoxes shqiperorum ) from the genus Rhabdodon and introduced the superordinate taxon of the Rhabdodontidae for both genera. The finds from Lower Austria were again listed in this publication under the name Mochlodon suessi . The authors considered it possible that they represented a further independent genus within the Rhabdodontidae, but left the question open for the time being with the reference to a re-evaluation of the relevant evidence that was currently in progress.
The by Weishampel et al. The revision mentioned in 2003 was only published in 2006. The authors of this study came to the conclusion that the supporting material was insufficient for an exact diagnosis and interpreted the Lower Austrian finds as a young animal of a species of the genus Zalmoxes ( Zalmoxes sp. ) Which cannot be identified . At the same time it was pointed out that the two vertebral bodies mentioned by Bunzel in his first description could no longer be found.
In 2012 the matter took a dramatic turn. Attila Ősi and several co-authors, including Weishampel, reported extensive new evidence from the Csehbánya formation of the Iharkút bauxite open- cast mine in Veszprém County in Hungary, which in terms of anatomical details and size largely matched the findings of Muthmannsdorf. The genus Mochlodon was reintroduced with Mochlodon suessi as a type species as a valid independent taxon and supplemented by a second species Mochlodon vorosi from the Hungarian site. The additional species “ vorosi ” honors the Hungarian paleontologist Attila Vörös .
Fossil evidence and age classification of the finds
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Mochlodon suessi (type species)
Fossil finds of this kind are only known from the historical site near Muthmannsdorf. According to Ősi et al., 2012, the supporting material includes:
- The fragment of a right lower jaw with several teeth (PIUW 2349/2; lectotype for type species and genus)
- A single, isolated tooth of the lower jaw (PIUW 2349/3)
- A single, isolated tooth of the upper jaw (PIUW 2349/4)
- The fragment of a parietal bone (PIUW 2349/54; originally identified by Bunzel as the parietal bone of a lizard)
- An incomplete left shoulder blade (PIUW 3518)
- A spoke fragment in question (PIUW 3517)
- A single, isolated claw bone , presumably from one of the front limbs (PIUW 2349/38)
- The fragment of a left femur (PIUW 2349 / III)
- The fragment of a presumably right tibia (PIUW 2348/35)
All specimens come from the Grünbach Formation as part of the Gosau Group and are kept at the Paleontological Institute of the University of Vienna (PIUW). The coal-bearing Grünbach Formation can be assigned to the lower Campanium on the basis of biostratigraphically usable fossil finds from the underlying Maiersdorf Formation and the overlying Piesting Formation , which corresponds to an age of about 83.5-76 Ma .
Mochlodon vorosi
There is much more evidence of the second type:
- A complete left lower jaw with four broken teeth (MTM V 2010.105.1; holotype of the species)
- Four more, almost complete lower jaws, two left (MTM V 2010.105.1, MTM 2012.15.1) and two right (MTM V 2010.107.1, MTM V 2010.109.1), and a total of six lower jaw fragments
- A left postorbital (MTM 01/14/2012)
- Two right quadrata (MTM V 2010.110.1, MTM V 2010.111.1)
- A total of 15 individual teeth in the upper jaw and 23 individual teeth in the lower jaw
- Four individual vertebrae from different sections of the spine
- An almost complete, but flattened sacrum (MTM V 2010.121.1)
- A total of three raven legs (MTM V 01.53, MTM V 2010.122.1, MTM V 2010.123.1) and a fragmentary shoulder blade (MTM 2012.22.1)
- A complete humerus (MTM V 2010.128.1) and a fragment of another (MTM 2012.23.1)
- A complete ulna (MTM 2012.24.1)
- Two almost complete (MTM V 01.225, MTM V 2010.126.1) and a fragmentary thighbone (MTM 2012.25.1)
- One complete tibia (MTM V 2010.127.1) and two tibial fragments (MTM V 01.101, MTM 2012.26.1)
- Two phalanges (MTM 2012.27.1, MTM 2012.28.1)
All fossil records come from the Csehbánya Formation and are kept at the Hungarian Natural Science Museum (Magyar Természettudományi Múzeum - MTM) in Budapest . The Csehbánya formation can be placed in the santonium (about 86.3–83.5 mya ) on the basis of palynological findings . Mochlodon vorosi is therefore not only older than Mochlodon suessi , but at the same time also the oldest known representative that can be safely assigned to the Rhabdodontidae.
features
Height and mass
Mochlodon is a genus of relatively small- stature representatives of the Iguanodontia. On the basis of the existing fossil record, the total body length of the adult animals can be determined to be about 1.6–1.8 m. Mochlodon was thus significantly smaller than the closely related genus Zalmoxes , for which a maximum total body length of up to 2.5 m is likely. The genera Mochlodon and Zalmoxes from Austria, Hungary and Romania are in clear contrast to the genus Rhabdodon from France and Spain, which could reach a total body length of up to 6 m.
A body mass of 31 kg is estimated for Mochlodon vorosi .
Generic and species-typical characteristics
(Abbreviations in brackets refer to the illustration on the right. The description of the generic features is based on the description by Ősi et al., 2012.)
The genus Mochlodon differs from other representatives of the Rhabdodontidae in particular through some features of the toothed lower jaw (dentals):
The outside of the lower jaw shows a shallow depression (“dep”) caudally , immediately below the coronoid process (“cop”). This feature is in Mochlodon suessi pronounced weaker than vorosi Mochlodon . The more extensive evidence for Mochlodon vorosi also suggests that the trait was more pronounced in larger individuals than in smaller ones. The surface of this depression is sculptured with fine, rustrocaudal oriented bone ridges. The physiological function of this depression has not yet been conclusively clarified, but it is suspected that it is a reinforced attachment point for the external jaw adductors .
The symphysis (“sy”), the contact surface of the cartilaginous connection between the two branches of the lower jaw, is only slightly curved inward. The dorsal edge (“ded”) of the symphysis is straight, parallel to the longitudinal axis of the lower jaw ( Mochlodon vorosi ) or is only slightly rostroventrally inclined ( Mochlodon suessi ). In this area there is also a deep pit (“gr”) on the upper edge of the lower jaw, which widens caudally.
Mochlodon vorosi also differs from Rhabdodon and Zalmoxes in the shape of the square leg , the head (“qh”) of which is strongly curved backwards. In Mochlodon suessi there is no corresponding fossil evidence for this element of the primary TMJ .
Dentition
(Abbreviations in brackets refer to the figure on the right. The description of the teeth is based on the descriptions by Ősi et al., 2012 and Virág & Ősi, 2017.)
As with other representatives of the Rhabdodontidae, the teeth of the upper and lower jaw differ in shape:
The crown of the maxillary teeth has an approximately rounded rectangular outline in the labial and lingual view. The wear area ("wf") is lingual. The tooth enamel (“en”) on the labial side is ornamented with 8–13 approximately parallel and equally strong longitudinal ribs (“lr”), which are framed by a slightly raised edge (“cr”) at the base of the crown as well as mesially and distally . The mesial and distal edges of the crown are slightly serrated. In the case of less worn upper jaw teeth, the enamel on the lingual side also shows an ornamentation with weakly pronounced, parallel longitudinal ribs, which however disappear with increasing wear.
The crown of the mandibular teeth has an approximately diamond-shaped outline in the labial and lingual view. The wear surface is labial. In contrast to the teeth of the upper jaw, the enamel on the lingual side is divided into two U-shaped fields by a particularly pronounced, centrally located, primary longitudinal rib (“pr”). The primary longitudinal rib is accompanied on each of these fields by 5–7 less pronounced secondary ribs ("sr"), which as a rule do not reach to the base of the crown. The mesial and distal edges of the crown are slightly serrated (“dm”). A slightly raised edge at the base of the crown is missing. The labial side of less worn lower jaw teeth shows, similar to the lingual side of the upper jaw teeth, an ornamentation with weakly pronounced, parallel longitudinal ribs, which disappears with increasing wear.
The simple roots of the upper jaw teeth are robust and reach a length that corresponds to about three times the crown length, while the roots of the lower jaw teeth only reach a maximum of about 1.5 times the crown length. In both cases there are flat, elliptical indentations ("rgr") on the lingual side of the roots, which correspond to the position of the crown of the replacement tooth that is pushing in.
The worn surfaces of the teeth show straight, approximately parallel scratches, which suggest a largely local chewing movement of the jaw. This means that the upper and lower jaws only moved vertically, but not palinally (back and forth) or transversely (sideways) to one another. A few individual teeth, which were originally determined to be lower jaw teeth by Mochlodon vorosi , show a different wear pattern with almost concentrically arranged, arched scratches, which suggest a simultaneous combination of orthal and palinal jaw movements during the chewing process. Such a circumpalinale chewing movement is unknown by representatives of the ornithopods, but has been found in some representatives of the Neoceratopsia , whose teeth may be very similar to those of ornithopods. It is therefore assumed that the individual teeth with different wear patterns are not to be assigned to Mochlodon vorosi , but to Ajkaceratops kozmai , the only known representative of the Neoceratopsida from the Csehbánya Formation.
Systematics
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| Systematic position of the genus Mochlodon within the Ornithopoda, simplified from Herne et al., 2019. |
The adjacent cladogram shows the systematic position of Mochlodon within the ornithopoda, simplified from Matthew C. Herne et al., 2019.
In this phylogenetic analysis, Mochlodon is the sister taxon of Zalmoxes . The two small genera of Eastern Europe form their own subclade within the Rhabdodontidae, which is opposed to the larger, Western European genus Rhabdodon . According to the age of Mochlodon vorosi , the division into small, Eastern European and large, Western European forms must have taken place before the Santonium.
The Rhabdodontidae themselves form an independent subclade within the group of Iguanodontia, which is opposed to all other representatives of this group. They are roughly equivalent in rank to a family and are endemic to Europe .
Weishampel et al. had originally defined the Rhabdodontidae as "the youngest common ancestor of Zalmoxes robustus and Rhabdodon priscus and all of its descendants" (knot-based taxon). However, this definition was later reformulated in the sense of the general suggestions of Paul Sereno to "all representatives of the Iguanodontia that are more closely related to Rhabdodon priscus than to Parasaurolophus walkeri " or, equivalent, but in other words: "the most comprehensive clade containing Rhabdodon priscus , but not Parasaurolophus walkeri ”(stem-based taxon).
In 2016, Paul-Emile Dieudonné and co-authors identified the Australian genus Muttaburrasaurus as the sister taxon of the Rhabdodontidae and introduced the parent taxon of the Rhabdodontomorpha (Rhabdodontidae + Muttaburrasaurus ; knot-based taxon). In 2019, however, Herne et al. Contradicted this interpretation, who in their analysis found Muttaburrasaurus in a common clade with the Dryomorpha , i.e. apart from the Rhabdodontidae.
Paleecology
During the Upper Cretaceous Period, Central Europe consisted of a collection of larger and smaller, largely isolated islands ("European Cretaceous Archipelago").
The sediments of the coal-bearing Grünbach Formation were deposited on the coastal fringes and the adjacent, further inland areas of one of these islands. Numerous plant fossils offer insight into the complex ecosystem . Ponds and backwaters were at least partially overgrown with aquatic plants of the genus Quereuxia and Brasenites . The flora of the adjacent wetlands was of ferns and pandanus plants ( Pandanites dominated). Marsh , breakage and swamp forests were representatives of the walnuts , palm ( Sabalites ) and evergreen coniferous trees of the genus Geinitzia constructed. Ferns of the genus Sphenopteris and trees and shrubs from the plane tree family grew on the banks of streams and rivers , the latter with Ettingshausenia forming the dominant floral element of this location. Mesophytic forests in somewhat higher and drier locations were dominated by conifers of the genus Pagiophyllum . The paleoflora of the Grünbach formation suggests a humid , subtropical climate with hot summers and relatively short, non- arid dry seasons and frost-free winters.
The alluvial sediments of older Csehbánya lineup was one of the larger of these islands in a lowland hem alluvial plain deposited. Further towards the coast, the alluvial plains merged into the freshwater swamps and lakes of the Ajka coal formation. The pollen spectrum of the Csehbánya Formation is dominated by representatives of the Normapolles group (40%), which presumably come from plants related to the beech-like . Pollen from other angiosperms contribute a further 15% and spores from mosses and ferns together make up 37%, while pollen from gymnosperms only play a minor role with 8%. The alluvial plain of the Csehbánya Formation was probably overgrown with Normapolles dominated mixed deciduous forests. Often there are nuts of the genus Sphaeracostata , which are assigned to this group. Magnolias of the genus Padragkutia and conifers of the araucaria family are subordinate elements of these forests, with the latter also having the possibility that their remains were washed into the storage room from higher areas of the hinterland. The undergrowth was made up of ferns and herbaceous flowering plants. Local accumulations of nettles are seen as a sign of a moderately damp subsoil and frequent disturbances of the ecosystem. The bank areas of shallow lakes and ponds were shaped by populations of Pandanites . In sum, the paleoflora of the Csehbánya formation also suggests a humid, subtropical climate with seasonal dry periods.
Mochlodon was, like all representatives of the Rhabdodontidae, probably a pure herbivore . The scissor-like closing dentition was well suited to cut to relatively hartfaserige plant parts and some authors suggest that the Rhabdodontidae generally to corresponding monocots plants such Sabalites or Pandanites were specialized.
At the beginning of the 20th century, Franz von Nopcsa had already started considering whether the small, Upper Cretaceous ornithopods from Austria and Romania might be a case of island dwarfing . On the basis of the findings by Mochlodon vorosi , Ősi et al., 2012, however, put the alternative hypothesis into the space that the genera Mochlodon and Zalmoxes do not represent cases of island dwarfing, but on the contrary, the large genus Rhabdodon represents a case of island gigantism .
Individual evidence
- ↑ A. Gaudry: Les du monde animal dans les temps Enchaînements géologiques: Fossil secondaires. Volume 3, Paris, 1890, p. 222, ( digitized version ).
- ^ A b c d E. Bunzel: The reptile fauna of the Gosau Formation in the New World near Wiener Neustadt. In: Treatises of the k. & k. Geological Reichsanstalt , Volume 5, Number 1, 1871, 18 pages + 8 plates, ( digitized version ).
- ↑ B. Plöchinger, G. Bardossy, R. Oberhauser & A. Papp: The Gosau basin of Grünbach and the New World (Lower Austria). In: Jahrbuch der Geologische Bundesanstalt , Volume 104, 1961, pp. 359-441, ( digitized version ).
- ↑ a b c d e f g h i j k l m n A. Ősi, E. Prondvai, R. Butler & DB Weishampel: Phylogeny, Histology and Inferred Body Size Evolution in a New Rhabdodontid Dinosaur from the Late Cretaceous of Hungary. In: PLoS ONE , Volume 7, Number 9, 2012, e44318, doi : 10.1371 / journal.pone.0044318 .
- ↑ a b HG Seeley: The Reptile Fauna of the Gosau Formation preserved in the Geological Museum of the University of Vienna. With a Note on the Geological Horizon of the Fossils at Neue Welt, west of Wiener Neustadt. In: The Quarterly Journal of the Geological Society of London , Volume 37, 1881, pp. 620-707, ( digitized ).
- ↑ F. Nopcsa: Dinosaur remains from Transylvania (skull of Limnosaurus transsylvanicus nov. Gen. Et spec.). In: Memoranda of the Imperial Academy of Sciences Vienna, Mathematisch-Naturwissenschaftliche Classe , Volume 68, 1900, pp. 555-591, ( digitized version ).
- ^ F. Nopcsa: Dinosaur remains from Transylvania II (skull remains from Mochlodon) with an appendix: on the phylogeny of the ornithopodids. In: Memoranda of the Imperial Academy of Sciences Vienna, Mathematisch-Naturwissenschaftliche Classe , Volume 72, 1902, pp. 149–175, ( digitized version ).
- ^ F. Nopcsa: The dinosaurs of the Transylvanian parts of Hungary. In: Communications from the yearbook of the Royal Hungarian Geological Institute , Volume 23, Number 1, 1915, pp. 1–24, ( digitized version ).
- ^ F. Nopcsa: On the Geological Importance of the Primitive Reptilian Fauna of the Uppermost Cretaceous of Hungary; with a description of a New Tortoise (Kallokibotium). In: Quarterly Journal of the Geological Society of London , Volume 79, 1923, pp. 100-116, ( digitized ).
- ^ F. Nopcsa: Sexual Differences in Ornithopodous Dinosaurs. In: Palaeobiologica , Volume 2, 1929, pp. 187-200, ( digitized version ).
- ^ A b W. Brinkmann: Rhabdodon Matheron, 1869 (Reptilia, Ornithischia): proposed conservation by suppression of Rhabdodon Fleishmann, 1831 (Reptilia, Serpentes) - ZN (S.) 2536. In: The Bulletin of Zoological Nomenclature , Volume 43, Number 3, 1986, pp. 269-272, ( digitized version ).
- ↑ ICZN: Opinion 1483 - Rhabdodon Matheron, 1869 (Reptilia, Ornithischia): conserved. In: The Bulletin of Zoological Nomenclature , Volume 45, Number 1, 1988, pp. 85-86, ( digitized ).
- ↑ a b DB Weishampel, C.-M. Jianu, Z. Csiki & DB Norman: Osteology and phylogeny of Zalmoxes (ng), an unusual Euornithopod dinosaur from the latest Cretaceous of Romania. In: Journal of Systematic Palaentology , Volume 1, Number 2, 2003, pp. 65-123, ( digitized version ).
- ↑ S. Sachs & JJ Hornung: Juvenile ornithopod (Dinosauria: Rhabdodontidae) remains from the Upper Cretaceous (Lower Campanian, Gosau Group) of Muthmannsdorf (Lower Austria). In: Geobios , Volume 39, Number 3, 2006, pp. 415-425, ( summary ).
- ↑ Gerald Hofer, Erich Draganits, Michael Wagreich, Christa-Charlotte Hofmann, Doris Reischenbacher, Marie-Louise Grundtner & Magda Bottig: Stratigraphy and Geochemical Characterization of Upper Cretaceous Nonmarine - Marine Cycles (Grünbach Formation, Gosau Group, Austria). In: Austrian Journal of Earth Sciences , Volume 104, Number 2, 2011, pp. 90-107, ( digitized version )
- ^ A b ER Bodor & V. Baranyi: Palynomorphs of the Normapolles group and related plant mesofossils from the Iharkút vertebrate site, Bakony Mountains (Hungary). In: Central European Geology , Volume 55, Number 3, 2012, pp. 259-292, ( digitized version ).
- ↑ RBJ Benson, NE Campione, MT Carrano, Ph. D. Mannion, C. Sullivan, P. Upchurch & DC Evans: Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage. In: PLoS Biology , Volume 12, Number 5, 2014, e1001853, doi : 10.1371 / journal.pbio.1001853 .
- ↑ a b A. Virág & A. Ősi: Morphometry, Microstructure, and Wear Pattern of Neornithischian Dinosaur Teeth From the Upper Cretaceous Iharkút Locality (Hungary). In: The Anatomical Record - Advances in Integrative Anatomy and Evolutionary Biology , Volume 300, Number 8, 2017, pp. 1439–1463, doi : 10.1002 / ar.23592 .
- ↑ FJ Varriale: Dental microwear reveals mammal-like chewing in the neoceratopsian dinosaur Leptoceratops gracilis. In: PeerJ , Volume 4, 2016, e2132, doi : 10.7717 / peerj.2132 .
- ↑ a b c d Matthew C. Herne, JP Nair, AR Evans & AM Tait: New small-bodied ornithopods (Dinosauria, Neornithischia) from the Early Cretaceous Wonthaggi Formation (Strzelecki Group) of the Australian-Antarctic rift system, with revision of Qantassaurus intrepidus Rich and Vickers-Rich, 1999. In: Journal of Paleontology , Volume 93, Number 3, 2019, pp. 543-584, ( digitized ).
- ↑ PC Sereno: The Logical Basis of Phylogenetic Taxonomy. In: Systematic Biology , Volume 54, Number 4, 2005, pp. 595-619, ( digitized ).
- ^ A b P. Godefroit, G. Garcia, B. Gomez, K. Stein, A. Cincotta, U. Lefèvre & X. Valentin: Extreme tooth enlargement in a new Late Cretaceous rhabdodontid dinosaur from Southern France. In: Nature - Scientific Reports , Volume 7, 2017, Article Number 13098, doi : 10.1038 / s41598-017-13160-2 .
- ↑ a b P.-E. Dieudonné, Th. Tortosa, FT Fernández-Baldor, JI Canudo & I. Díaz-Martínez: An Unexpected Early Rhabdodontid from Europe (Lower Cretaceous of Salas de los Infantes, Burgos Province, Spain) and a Re-Examination of Basal Iguanodontian Relationships. In: PLoS ONE , Volume 11, Number 6, 2016, e0156251, doi : 10.1371 / journal.pone.0156251 .
- ↑ a b Z. Csiki-Sava, E. Buffetaut, A. Ősi, X. Pereda-Suberbiola & St. L. Brusatte: Island life in the Cretaceous - faunal composition, biogeography, evolution, and extinction of land-living vertebrates on the Late Cretaceous European archipelago. In: ZooKeys , Volume 469, 2015, pp. 1-161, doi : 10.3897 / zookeys.469.8439 .
- ^ AB Herman & J. Kvaček: Late Cretaceous Grünbach Flora of Austria. Naturhistorisches Museum Wien, Vienna, 2010, ISBN 978-3-902421-43-2 , 224 pages, ( digitized version ).
- ↑ a b G. Botfalvai, J. Haas, ER Bodor, A. Mindszenty & A. Ősi: Facies architecture and palaeoenvironmental implications of the upper Cretaceous (Santonian) Csehbánya formation at the Iharkút vertebrate locality (Bakony Mountains, Northwestern Hungary) In: Palaeogeography, Palaeoclimatology, Palaeoecology , Volume 441, Number 4, 2016, pp. 659-678, ( abstract ); ( Manuscript version ).
- ↑ J. Kocsis, A. Ősi, T. Vennemann, CN Trueman & MR Palmer: Geochemical study of vertebrate fossils from the Upper Cretaceous (Santonian) Csehbánya Formation (Hungary): Evidence for a freshwater habitat of mosasaurs and pycnodont fish. In: Palaeogeography, Palaeoclimatology, Palaeoecology , Volume 280, 2009, pp. 532-542, ( abstract ).
