Strongly completed skeleton reconstruction of Megalosaurus
|Middle Jurassic ( Bathonium )|
|168.3 to 166.1 million years|
|Buckland , 1824|
Megalosaurus is a genus of large dinosaurs from the Theropoda group . She lived about 168 to 166 million years ago in the middle Jurassic of Western Europe . As the first ever scientifically named genus of dinosaurs, it has specialsignificance in the history of science . The genus includes only one generally recognized nominal species , the type species Megalosaurus bucklandii .
The generic name is a Latinized ancient Greek compound from the reinforcing prefix μέγαλο- mégalo- ' big , high, great, very' and σαῦρος sauros 'lizard'. It appears first in the Outlines of Oryctology by the British doctor James Parkinson , published in 1822 . However, the British geology and paleontology pioneer William Buckland (1824) is usually cited as the author of the name or as the first scientific descriptor of the genus . This is also honored by the epithet of the type species. The British doctor Gideon Mantell (1827) is considered to be the author of the binomial Megalosaurus bucklandii , whose name, like that of Buckland, is closely linked to the beginnings of dinosaur research (see also → Iguanodon ).
Material and sources
Find region of M. bucklandii (based on Benson, 2010)
The material known from the type species comes from several individuals and from different localities, but is limited to only a few, often not even completely traditional elements of the skeleton. It comprises (as of 2010) a few dermal bones of the skull (maxillaries, jugals, dentals and fragments of some posterior lower jaw bones), several isolated teeth, a few vertebrae, some ribs, the elements of the shoulder girdle, several humeri, just a single forearm bone (an ulna), a single finger distal phalanx, several sacrum and several pelvic bones, several thigh bones (femora), several lower leg bones (but only "shins", tibiae) and several metatarsals (metatarsals).
All material comes from deposits of the Central Jurassic Great Oolite Group of the Cotswolds in southern England . Stratum typicum or Locus typicus for M. bucklandii is the “Stonesfield Slate” facies of the Taynton Limestone Formation ( Bathonium ) of Stonesfield in West Oxfordshire . The remaining material comes from the Sharp's Hill Formation of Chipping Norton , West Oxfordshire, and from unspecified layers of the Great Oolite Group of Sarsgrove near Chipping Norton and from the New Park Quarry quarries in Longborough and Oakham Quarry near Little Compton (both in the Chipping Norton Limestone Formation) in east Gloucestershire . All deposits of the Great Oolite Group were deposited in a shallow marine environment, that is, the Megalosaurus remains were not embedded in the sediment where the animals lived, but were washed into the sea by rivers from one of the surrounding land areas and embedded there . This transport also explains the accumulation of certain skeletal elements (pelvic bones, long bones) in the known material and the extensive or complete absence of other elements (many skull bones, vertebrae, ribs, hand bones, phalanges of the toes).
In the most recent revision of the type species M. bucklandii (Benson, 2010) the following "unique combinations of features of the dental" (i.e. the tooth-bearing bone of the lower jaw) are named as diagnostic ( also diagnostic for the genus due to their monotype ): 13 to 14 tooth positions ; third tooth socket (third alveolus) not enlarged (in contrast to all other spinosauroids ) * ; Dentals in a view from above (dorsal) running straight, with non-enlarged symphysial region ; outside (lateral) row of passages for nerves and blood vessels (neurovascularforamina) lies in a shallow channel; Interdental plates high and not fused together; two Meckel's foramen; shallow Meckel channel.
Furthermore, Benson (2010) names the following aut apomorphies of the genus, which are not shown in the lectotype (a dental one), but in associated material outside the type series: ventral surfaces of sacral vertebrae 1 and 3 to 5 evenly rounded, ventral The surface of the 2nd sacral vertebra, however, has an angular longitudinal back; weak dorsally directed sine in the middle section of the dorsal margin of the shoulder blade; Bone bulge on the lateral surface of the ilium (median iliac ridge) with several parallel longitudinal furrows, which give the surface of the bulge a rippled appearance, obliquely from the acetabulum upwards (posterodorsally); the protrusions that run along the inside (medial side) of the shaft of the two ischia (English ischial aprons ) are thickened in the head-to-tail direction (cranio-caudal) (in contrast to the narrow "ischial aprons" of Allosaurus and Torvosaurus ) and have an approximately flat inner (medial) surface; the muscle attachment point ( distal ischial tubercle ) located approximately in the middle of the lateral side of the shaft of the ilium has a wrinkled surface; Complementary "groove-and-tenon" structures on the joint surfaces between metatarsals II and III.
Of all the other members of the megalosauridae differs Megalosaurus by a pneumatisiertes Jugale, spinous processes of the dorsal vertebrae, which are about twice as high as the vortex centers, and a tilted forward (craniad) Deltopectoralfortsatz (large muscle attachment point on the upper arm bone). From all other members of the Megalosauroidea (megalosauroidea) differs Megalosaurus by the non-enlarged third tooth socket in the dental * and in that the front branch of the maxilla is higher than long.
Although Megalosaurus is one of the longest known dinosaurs, it is one of the lesser-known genera of dinosaurs because of the relatively sparse remains that have been passed down of it. Everything that can be said about his appearance and physique must be inferred either from the material available or from better known, relatively closely related species. So part Megalosaurus , based on the size of the traditional bone, probably the largest theropod the Middle Jurassic: Adult animals reach a head-to-tail length of 8 to 9 meters and a body mass of probably more than 950 kilograms. The strong, sturdy bones suggest that Megalosaurus had a muscular, stocky build, but was probably not quite as sturdy as Torvosaurus . The straight dental line up to the symphysis suggests that the muzzle was relatively pointed. Otherwise he would have shown the typical appearance of a tetanur (that is, a relatively "primitive" theropod), with comparatively long arms and three-fingered, clawed hands.
Taxonomy and systematics
Prehistory and first description
The first mention and illustration of a bone fragment supposedly from Megalosaurus can be found in the work Natural History of Oxford-shire by the English scholar Robert Plot , published in 1677 . It was the lower (distal) end of a thigh bone (femur) with the two joint heads (condyli femoris) - the bone shaft (diaphysis) was missing. The piece, which according to the plot came from Cornwell (near Chipping Norton ), has been lost over the centuries and its exact taxonomic identity can therefore no longer be determined today. However, its size, appearance, and origin from an Oxfordshire quarry suggest it was likely at least a dinosaur femur. Plot speculated at the time that it either came from a Roman war elephant , or from a human being about three meters tall, although the latter interpretation seemed to him the more plausible. In 1763 the piece was reproduced by the English doctor and naturalist Richard Brookes in the 5th volume of his New and accurate system of natural history (as part of a largely pure reproduction of the representations Plots). Because of its external resemblance to a human scrotum , he labeled it Scrotum humanum , and it is not clear from his descriptions whether he was aware that this piece was the fossil, identified the plot as part of a femur and had described. **
In the further course of the 18th century, a number of putative and probable dinosaur bones, the whereabouts of which we know today, were excavated in Stonesfield. But William Bucklands (1824) and Gideon Mantells (1827) formal initial descriptions of the genus Megalosaurus and the species M. bucklandii were based in part on material that had been known for some time. Specifically, *** dentals, which were established as a lectotype long later, were discovered as early as 1797. A novelty in Buckland's description was that he interpreted the fossil bones from Stonesfield as the remains of a giant reptile. However, he was inspired by the French naturalist Georges Cuvier , who met Buckland in 1818 and examined the bones. Buckland's (future) wife Mary also had her share by making drawings and establishing contact with Cuvier through correspondence. Buckland and Mantell saw in Megalosaurus a kind of giant lizard and compared him to monitor lizards and crocodiles . Furthermore, Mantell in particular suspected that teeth and bones from several other regions and strata of England came from Megalosaurus . ¶ However, this was due to the at that time not yet existing knowledge of the diversity of theropods, the low specific diagnostic value of theropod teeth and the considerable numerical age differences between the respective layer sequences.
Further taxonomic history in the 19th and 20th centuries
In 1842 Richard Owen finally established the order Dinosauria, based on the three genera Iguanodon , Hylaeosaurus and Megalosaurus . Even if none of the characteristics that Owen considered relevant at the time (e.g. the five-vertebral sacrum) still has a diagnostic value for the group today, and some already known taxa (e.g. Plateosaurus , Cetiosaurus , Poekilopleuron ) have not yet been recognized as belonging were, the concept of the Dinosauria has been able to hold up to this day due to its initial connection with these three genera.
In the next 100 years or so, numerous Megalosaurus species from different regions of the world, mostly from Europe, were described. The finds on which these species were justified come from the entire dinosaur-bearing interval from the Upper Triassic to the Upper Cretaceous and mostly include isolated individual bones or teeth, which, according to current knowledge, cannot be determined at the genus level and in some cases cannot even be assigned to dinosaurs.
- For example, Friedrich August Quenstedt described the species Megalosaurus cloacinus in 1858 from Oberkeuper in Württemberg ("Rhät- Bonebed ") using isolated teeth. With him it says literally: “ The sickle-shaped teeth, which are sharp and finely notched on the concave side, but round and smooth below on the convex side, evidently speak for the sex [d. H. the genus Megalosaurus ]. ”Quenstedt's“ Keuper teeth ”agree in these characteristics with the teeth of the holotype of M. bucklandii , but not only with these, but also with the teeth of numerous other theropods known today. Therefore today they cannot be assigned to a specific genus and not even to a specific large group within the theropods.
- In 1870 Jean-Baptiste Greppin described the species M. meriani from the Upper Jura of the Canton of Bern using a tooth and a few damaged postcranial elements. The latter, however, were later largely recognized as the remains of sauropods and re-described under the name Ornithopsis (?) Greppini , so that only the tooth remained in the type material of M. meriani . This probably belongs to a ceratosaur .
- In the same year, Gustave Lennier described M. insignis on the basis of a single tooth without a point from the layer sequence of the lower Kimmeridgium (Upper Jurassic) at the Cap de La Hève in Le Havre . Today the piece can only be assigned to an indefinable theropod.
- In 1881 Harry Govier Seeley described M. pannoniensis on the basis of two individual teeth from the Upper Cretaceous of Austria ( Gosau layers ), which may come from either a dromaeosaurid or a tyrannosauroid .
- In 1902 Franz von Nopcsa described M. hungaricus on the basis of a single tooth from the " reptile-leading Cretacian coals from Nagy-Bároth " in Transylvania (now Romania), then part of Austria-Hungary . The tooth may belong to a dromaeosaurid or tyrannosauroid.
- In 1903, Louis named Dollo M. lonzeensis on the basis of an individual finger phalanx from the glauconite marl from Lonzée ( Coniac - Santon ) in the province of Namur , Belgium, which he himself had mentioned and described for the first time 20 years earlier . The piece probably comes from a coelurosaur .
- In 1921 Werner Janensch described the species M. ingens based on a single large tooth from the Upper Jurassic of the famous Tendaguru fossil deposit in Tanzania . This and other teeth from different stratigraphic levels of this site belong to non-determinable representatives of the (more basal) Tetanurae.
- 1926 renames Friedrich von Huene in a work in which he intensively with Megalosaurus apart sets and among other already relatively modern skulls and skeletal reconstruction for M. bucklandii delivers § the new kind M. lydekkeri . It is based on a single tooth from the lower Lias group (? Sinemurium ) from Lyme Regis in Dorset, which Richard Lydekker (1888) depicted as " Zanclodon (?) Sp." The piece later put under reserve by Huene in “his” genus Magnosaurus (see below) probably represents a relatively basal theropod outside the Tetanurae.
All these and other taxon names are now considered to be the nomina dubia . While Quenstedt's assignment of his "Keuper tooth" to Megalosaurus can still be excused with the generally low level of knowledge of his time, the concept of the genus developed more and more into one with the increase in knowledge about the diversity of theropods towards the end of the 19th century so-called trash can taxon. Nopcsa (1915) remarked: “ So [...] the name Megalosaurus is still today for the younger theropods a container into which one puts all the European theropod remains of the Jurassic and the Chalk that cannot be precisely determined. For the time being we have no means to get rid of this evil [...]. "
In addition to the historical species, which are based on undiagnostic material, several Megalosaurus species were described on the basis of diagnostic fossils, but later, due to clear differences from the type material of M. bucklandii, into separate, newly established genera , some of which are only relatively distantly related to Megalosaurus posed. The best known of these species include Majungasaurus crenatissimus ( Depéret , 1896) $ , an abelisaurid from the Upper Upper Cretaceous of Madagascar, Carcharodontosaurus saharicus ( Depéret & Savornin , 1925) § , an allosauroide from the Higher Lower Cretaceous of North Africa, and Dilophosaurus wetherilli ( Welles , 1954), a basal neotheropod from the Lower Jurassic Arizona. Huenes' genus Magnosaurus, established in 1932 and now widely recognized, is based on a species originally assigned to Megalosaurus with M. nethercombensis ( Huene , 1926) from the lower Central Jurassic of Dorset .
In addition, some names were published without an image of the associated fossils or an adequate description. These nouns nuda include the " Megalosaurus dapukaensis " and " M. tibetensis " finds reported from China in the 1980s .
The last Megalosaurus species formally valid in the 20th century and first described on the basis of diagnostic material was M. hesperis Waldman from the Inferior Oolite ( Bajocium ) of Dorset in 1974, i.e. from roughly the same stratigraphic interval and almost exactly the same area ( Sherborne ) such as Magnosaurus nethercombensis . The corresponding fossil specimens were originally assigned to the type species Megalosaurus bucklandii . In the 2000s, the type material of M. hesperis and M. bucklandii was examined again in detail, whereby the researcher came to the conclusion that the two forms differ even at the genus level. Consequently, the new genus Duriavenator was established in 2008 for M. hesperis . The genus Megalosaurus has been monotypical since then . The validity of the genus Megalosaurus was confirmed in the same year by the same editor after it had been questioned in 2002. However, the use of the name Megalosaurus has been restricted for the time being to the lectotype of Megalosaurus bucklandii . In 2010, after a more extensive examination of the theropod material from the Great Oolite Group, another revision took place including the assignment of the entire historical type material of Megalosaurus bucklandii to this species and therefore to the genus (see material and sources ).
In the pre-cladistic, more modern classification schemes of the dinosaurs, Megalosaurus was considered a representative of the Carnosauria in the traditional sense, that is, the large, heavily built representative of the theropods . Within the Carnosauria he was considered more "primitive" representatives and formed with other such representatives (formerly or now also in part to the genus Megalosaurus had been put) the family megalosauridae .
Even in the modern, cladistic-based classification scheme of Saurischia, the main features of which can be traced back to Jacques Gauthier (1986), Megalosaurus occupies a relatively basic position as a basic representative of the very inclusive clade Tetanurae . As a result of various recent cladistic analyzes, Megalosaurus appears either within a paraphyletic series of developments at the base of the Tetanurae or within a basal Tetanurae clade, the Spinosauroidea or Megalosauroidea. The problem here is the relatively sparse fossil record of both Megalosaurus and closely related taxa, which in the cladograms to this day leads to instability and / or a relatively poor resolution of relationships (so-called polytomies). Already Gauthier (1986) spoke of a "'megalosaur' problem" in this context.
This can also be seen in the results of two analyzes of the relationships of the Tetanurae, which were published in 2012 after the most recent revision of Megalosaurus bucklandii . ¥ Although Megalosaurus has a sister group relationship with Torvosaurus in both results , this clade in Carrano et al. (2012) found a sister group relationship with Duriavenator , while Rauhut et al. (2012) has a sister group relationship with a polytomy of Dubreuillosaurus and Afrovenator . These common clades, in turn, form the phylogenetically newly defined Megalosauridae with the most closely related taxa.
Closer relationships between Megalosaurus according to Carrano et al. (2012)
Close relationships between Megalosaurus according to Rauhut et al. (2012)
- Roger BJ Benson: A description of Megalosaurus bucklandii (Dinosauria: Theropoda) from the Bathonian of the UK and the relationships of Middle Jurassic theropods. Zoological Journal of the Linnean Society. Vol. 158, No. 4, 2010, pp. 882-935, doi: 10.1111 / j.1096-3642.2009.00569.x .
- EA Howlett, WJ Kennedy, HP Powell, HS Torrens: New light on the history of Megalosaurus, the great lizard of Stonesfield. Archives of Natural History. Vol. 44, No. 1, 2017, pp. 82-102, doi: 10.3366 / anh.2017.0416
- Wilhelm Gemoll: Greek-German school and hand dictionary. Reviewed and expanded by Karl Vretska. With an introduction to the history of language by Heinz Kronasser. 9th edition. Freytag et al., Munich et al. 1965.
- James Parkinson: Outlines of oryctology. An introduction to the study of fossil organic remains; especially those found in the British strata. London 1822 ( archive.org ), p. 289 .
- William Buckland: Notice on the Megalosaurus or great Fossil Lizard of Stonesfield. In: Transactions of the Geological Society of London. Series 2, Vol. 1, No. 2, 1824, pp. 390-396 ( BHL ).
- Gideon Mantell: Illustrations of the Geology of Sussex, Containing a General View of the Geological Relations of the South-Eastern Part of England, with Figures and Descriptions of the Fossils of Tilgate Forest. London 1827 ( GoogleBooks ), p. 67
- Julia J. Day, Paul M. Barrett: Material referred to Megalosaurus (Dinosauria: Theropoda) from the Middle Jurassic of Stonesfield, Oxfordshire, England: One taxon or two? Proceedings of the Geologists Association. Vol. 115, No. 4, 2004, pp. 359-366, doi: 10.1016 / S0016-7878 (04) 80015-4
- Roger BJ Benson: An assessment of variability in theropod dinosaur remains from the Bathonian (Middle Jurassic) of Stonesfield and New Park Quarry, UK, and taxonomic implications for Megalosaurus bucklandii and Iliosuchus incognitus. Paleontology. Vol. 52, No. 4, 2009, pp. 857-877, doi: 10.1111 / j.1475-4983.2009.00884.x , p. 858.
- For details on the paleogeography of Great Britain in the Middle Jurassic see: AJM Barron, GK Lott, JB Riding: Stratigraphical framework for the Middle Jurassic strata of Great Britain and the adjoining continental shelf. British Geological Survey Research Report RR / 11/06. Keyworth 2012 ( NORA )
- cf. Fig. 1 in: Roger BJ Benson, Paul M. Barrett, H. Philip Powell, David B. Norman: The taxonomic status of Megalosaurus bucklandii (Dinosauria, Theropoda) from the Middle Jurassic of Oxfordshire, UK. Paleontology. Vol. 51, No. 2, 2008, pp. 419-424, doi: 10.1111 / j.1475-4983.2008.00751.x , p. 421.
- Matthew T. Carrano, Roger BJ Benson, Scott D. Sampson: The phylogeny of Tetanurae (Dinosauria: Theropoda). Journal of Systematic Palaeontology. Vol. 10, No. 2, 2012, pp. 211-300, doi: 10.1080 / 14772019.2011.630927 , supplementary file No. 2 (description of characteristics).
- Robert Plot: The natural History of Oxford-shire. Being an Essay toward the Natural History of England. Oxford 1677, doi: 10.5962 / bhl.title.23488 , p. 131 ff. , Plate VIII, Fig. 4 .
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- see e.g. B. Oliver Gerke, Oliver Wings: Multivariate and cladistic analyzes of isolated teeth reveal sympatry of theropod dinosaurs in the Late Jurassic of Northern Germany. PLoS ONE. Vol. 11, No. 7, 2016, Art-No. e0158334, doi: 10.1371 / journal.pone.0158334
- Ralph E. Molnar, Sergei M. Kurzanov, Dong Zhiming: Carnosauria. Pp. 169–209 in: David B. Weishampel, Peter Dodson, Halszka Osmólska (eds.): The Dinosauria. University of California Press, Berkeley 1990, pp. 203 f.
- Michael J. Benton, PS Spencer: Fossil Reptiles of Great Britain. Springer, Dordrecht 1995, ISBN 978-94-010-4231-4 , pp. 140-146 (subsection on the stone quarries of Stonesfield)
- Oliver Rauhut: The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD Thesis, University of Bristol, 2000
- Ronan Allain, Daniel J. Chure: Poekilopleuron bucklandii , the theropod dinosaur from the Middle Jurassic (Bathonian) of Normandy. Paleontology. Vol. 45, No. 6, 2002, pp. 1107-1121, doi: 10.1111 / 1475-4983.00277 .
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- Friedrich August Quenstedt: The Jura. Tübingen 1858 ( MDZ reader ), p. 33
- Matthew T. Carrano, Roger BJ Benson, Scott D. Sampson: The phylogeny of Tetanurae (Dinosauria: Theropoda). In: Journal of Systematic Palaeontology. Vol. 10, No. 2, 2012, pp. 211-300, doi: 10.1080 / 14772019.2011.630927 , pp. 257 ff.
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- Roger BJ Benson: The osteology of Magnosaurus nethercombensis (Dinosauria, Theropoda) from the Bajocian (Middle Jurassic) of the United Kingdom and a re-examination of the oldest records of tetanurans. Journal of Systematic Palaeontology. Vol. 8, No. 1, 2010, pp. 131-146, doi: 10.1080 / 14772011003603515
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- Opinion 2269 (Case 3487) - Megalosaurus crenatissimus Depéret, 1896 (currently Majungasaurus crenatissimus ; Dinosauria, Theropoda): designation of a neotype. The Bulletin of Zoological Nomenclature. Vol. 68, No. 1, 2011, pp. 89-90, doi: 10.21805 / bzn.v68i1.a7
- Stephen L. Brusatte, Paul C. Sereno: A new species of Carcharodontosaurus (Dinosauria: Theropoda) from the Cenomanian of Niger and a revision of the genus. Journal of Vertebrate Paleontology. Vol. 27, No. 4, 2007, pp. 902-916, doi : 10.1671 / 0272-4634 (2007) 27 [902: ANSOCD] 2.0.CO; 2
- cf. Michael J. Benton, PS Spencer: Fossil Reptiles of Great Britain. Springer, Dordrecht 1995, ISBN 978-94-010-4231-4 , p. 125
- Michael Waldman: Megalosaurids from the Bajocian (Middle Jurassic) of Dorset. Paleontology. Vol. 17, No. 2, 1974, pp. 325–339 ( palass.org )
- Roger BJ Benson: A redescription of ' Megalosaurus ' hesperis (Dinosauria, Theropoda) from the Inferior Oolite (Bajocian, Middle Jurassic) of Dorset, United Kingdom. Zootaxa. Vol. 1931, No. 1, 2008, pp. 57-67, doi: 10.11646 / zootaxa.1931.1.5
- Roger BJ Benson, Paul M. Barrett, H. Philip Powell, David B. Norman: The taxonomic status of Megalosaurus bucklandii (Dinosauria, Theropoda) from the Middle Jurassic of Oxfordshire, UK. Paleontology. Vol. 51, No. 2, 2008, pp. 419-424, doi: 10.1111 / j.1475-4983.2008.00751.x .
- Zhao Xijin: Phylogeny and evolutionary stages of Dinosauria. Acta Palaeontologica Polonica. Vol. 28, No. 1-2, 1983, pp. 295-306 ( app.pan.pl ), p. 302
- Jacques Gauthier: Saurischian monophyly and the origin of birds. Pp. 1-55 in: Kevin Padian (Ed.): The Origin of Birds and the Evolution of Flight. Memoirs of the California Academy of Sciences, Vol. 8. California Academy of Sciences, San Francisco 1986 ( BHL )
- see e.g. B. Thomas R. Holtz, Jr .: A new phylogeny of the carnivorous dinosaurs. In: Gaia - Revista de Geociências do museu Universidade Lisboa. Vol. 15, 1998, pp. 5-61.
- Matthew T. Carrano, Roger BJ Benson, Scott D. Sampson: The phylogeny of Tetanurae (Dinosauria: Theropoda). In: Journal of Systematic Palaeontology. Vol. 10, No. 2, 2012, pp. 211-300, doi: 10.1080 / 14772019.2011.630927 , pp. 247 ff.
- Oliver WM Rauhut, Christian Foth, Helmut Tischlinger, Mark A. Norell: Exceptionally preserved juvenile megalosauroid theropod dinosaur with filamentous integument from the Late Jurassic of Germany. In: Proceedings of the National Academy of Sciences. Vol. 109, No. 29, 2012, pp. 11746–11751, doi: 10.1073 / pnas.1203238109
- Jacques Gauthier: Saurischian monophyly and the origin of birds. Pp. 1-55 in: Kevin Padian (Ed.): The Origin of Birds and the Evolution of Flight. Memoirs of the California Academy of Sciences, Vol. 8. California Academy of Sciences, San Francisco 1986 ( BHL ), p. 10
- see e.g. B. Thomas R. Holtz, Jr., Ralph E. Molnar, Philip J. Currie: Basal Tetanurae. Pp. 71–110 in: David B. Weishampel, Peter Dodson, Halszka Osmólska (eds.): The Dinosauria. 2nd Edition. University of California Press, Berkeley 2004 p. 96