Campylognathoides ("curved jaw", Strand 1928) is a genus of pterosaur, discovered in the Württemberg Lias deposits (dated to the early Toarcian age[1]) of Germany; this first specimen consisted however only of wing fragments. Further better preserved specimens were found in the Holzmaden shale: basing on these specimens Felix Plieninger erected a new genus.[2]

Temporal range: Early Jurassic, 182 Ma
Campylognathoides liasicus cast - University of California Museum of Paleontology - Berkeley, CA - DSC04685
Cast of the Pittsburgh specimen of C. liasicus
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Order: Pterosauria
Clade: Novialoidea
Superfamily: Campylognathoidea
Kuhn, 1967
Family: Campylognathoididae
Kuhn, 1967
Subfamily: Campylognathoidinae
Kuhn, 1967
Genus: Campylognathoides
Strand, 1928
Type species
Campylognathus zitteli
Plieninger, 1894
  • C. zitteli (Plieninger, 1894)
  • C. indicus? Jain, 1978
  • C. liasicus (Quenstedt, 1858)


Campylogn DB

Compared to its contemporary from the same layers Dorygnathus, the snout on this genus is relatively short, though the skull is still in general elongated, be it much lighter built. The large eye sockets, placed low in the skull above a narrow jugal, have caused some researchers to speculate that Campylognathoides had especially acute vision, or possibly even a nocturnal lifestyle. The back of the skull is relatively high and flat, with a sudden downturn just in front of the eyes. The snout ends in a slender point curving a bit upwards at its very end. A large part of the snout is occupied by long bony nares. Below them a small triangular skull opening, the fenestra antorbitalis is present.

Reflecting the more shallow snout, the teeth of Campylognathoides are also short and not at all laniaries or fang-like as in the markedly heterodont Dorygnathus. They are conical and recurved but have a broad base with the point bevelled off from the inside forming a sharp and strong cutting surface. In the upper jaw there are four rather widely spaced teeth in the praemaxilla gradually increasing in size from the front to the back; the fourth pair of teeth is the largest. Behind them are ten smaller teeth in the maxilla, gradually decreasing posteriorely. In the lower jaw there are twelve to fourteen teeth present in C. liasicus, sixteen to nineteen in C. zitteli. The largest total number is thus 66.

According to a study by Kevin Padian there are eight cervical vertebrae, fourteen dorsals, four or five sacrals and up to 38 caudal vertebrae. The tail base is flexible with about six short vertebrae; behind them the caudals elongate and are stiffened by very long extensions allowing the tail to function as a rudder.[3]

The sternum of Campylognathoides was a rather large rectangular plate of bone with a short forward-facing crest called a cristospina.[2] The upper arm is short but robust with a square deltopectoral crest. The lower arm too is short but wing length is considerable due to the hand, which has short metacarpals but a very long wing finger for a basal pterosaur, of which the second phalanx is the largest. The pteroid is short and robust.

The pelvis is not very well known. A fossil collector found a well preserved Campylognathoides hip in a Braunschweig shale quarry in 1986. This pelvis, BSP 1985 I 87, proved to be scientifically significant because the hip socket was according to Peter Wellnhofer in an upward lateral position, preventing the animal from being able to orient its legs erectly like in dinosaurs, birds and mammals. This would prove that Campylognathoides was not well able to walk on its hind legs but must have walked quadrupedally.[4] This gait posture has been confirmed in other "rhamphorhynchoids" (i.e. basal pterosaurs) as well.[2] However, Padian in 2009 concluded the opposite, stating that an erected position was necessary to place the feet on the ground and that, though a quadrupedal gait was possible, a bipedal way of locomotion was a precondition for a fast gait.[5] This subject remains highly controversial.

The leg is rather short and the feet are small. The fifth toe, often interpreted as carrying a membrane between the legs, is exceptionally short for a basal pterosaur.


Campylognathoides sp AMNH 1713 cast skull
The skull of C. liasicus, part of a cast of a skeleton in the American Museum of Natural History

In 1858 Friedrich August Quenstedt named a new species of Pterodactylus: P. liasicus. It was based on a fossil, holotype GPIT 9533, consisting of some wing bones, found on the Wittberg near Metzingen in layers dating from the early Toarcian, about 180 million years old. The specific name referred to the Lias. Quenstedt thought he had identified long metacarpals in the wing, concluding that the new species was therefore not belonging to more basal genera, like the long-tailed Rhamphorhynchus.[6]

In 1893 commercial fossil collector Bernhard Hauff sr. discovered a skeleton of a large pterosaur near Holzmaden. In 1894 Felix Plieninger based a new genus on this specimen: Campylognathus. The genus name is derived from Greek kampylos, "bent", and gnathos, "jaw", in reference to the bent lower jaw. The type species is Campylognathus zitteli. The specific name honours Alfred von Zittel.[7] The holotype is SMNS 9787.

In 1897 Hauff prepared another specimen that eventually in 1903 was acquired by the Carnegie Museum of Natural History at Pittsburgh. This fossil, CM 11424, is due to its completeness the best source of information about the genus.

In 1901 Plieninger for the first time studied P. liasicus and discovered that Quenstedt had mistaken the, in reality short, metacarpal, for a coracoid, meaning it was a basal pterosaur.[8] In 1906 Plieninger referred P. liasicus and the Pittsburgh specimen to Campylognathus, though not yet establishing the specific status of each of the three exemplars.[9] In 1907 however, Plieninger recognised a second species of Campylognathus: C. liasicus, to which CM 11424 was referred also.[10]

Norwegian entomologist Embrik Strand discovered in the 1920s that the name Campylognathus had previously been used for the African bug Campylognathus nigrensis, a genus of the Heteroptera named in 1890. As the name was thus preoccupied, he renamed the pterosaur Campylognathoides in 1928.[11]

During the twentieth century new finds have brought the number of known specimens to about a dozen.


Three species of Campylognathoides have been named:

  • Campylognathoides zitteli (Plieninger, 1894) is the type species.
The larger Holzmaden Campylognathoides, C. zitteli, had a six foot (1.825 metres for SMNS 9787) wingspan.[2]
  • Campylognathoides liasicus (Quenstedt, 1858 [originally Pterodactylus liasicus])
C. liasicus had a three foot wingspan, making it smaller than its Holzmaden contemporary, C. zitteli.[2]
  • "Campylognathoides" indicus Jain, 1974
C. indicus was described by Sohan Lal Jain on the basis of a fragment of jaw, ISI R. 48, recovered from Chanda district, India.[12] Kevin Padian considers this a nomen dubium, possibly based on a fish fossil. That the Kota Formation in which it was found, has since been redated to the Middle Jurassic or later, seems to preclude any close connection to Campylognathoides, even if it were a pterosaur.[13]

The distinction between C. liasicus and C. zitteli is problematical. Plieninger merely recognised the smaller species because he considered its fossil too poor in quality to refer other specimens to. However, in 1925 Swedish researcher Carl Wiman, studying specimen UUPM R157, concluded that a fundamental morphological difference could distinguish the two species: C. zitteli has a proportionally much longer wing. In 2008 however, Padian pointed out that this might well have been a matter of ontogenetic development, larger individuals growing extra large wings to limit the wing load. Other differences, such as the larger number of teeth in the lower jaw, a longer snout and nares, five instead of four sacrals, perpendicular sacral ribs and a longer leg, might conceivably also be size-related. Final proof could only be given by a continuous growth series, as previously has been done in the case of Rhamphorhynchus and Pterodactylus. Provisionally Padian kept distinguishing two species, but moved two specimens to C. zitteli: SMNS 51100 and GPIT 24470, because of their larger size and morphological similarities.[14]


Skeletal restoration of C. liasicus

Plieninger in his later publications assigned Campylognathus to the "Rhamphorhynchoidea". As this suborder is a paraphyletic assemblage of not specially related basal pterosaurs, this classification merely states the negative fact that it was not a pterodactyloid. A positive determination was first attempted by Baron Franz Nopcsa who in 1928 assigned the genus to the subfamily Rhamphorhynchinae within the family Rhamphorhynchidae.[15] After a period in which very little work was done on pterosaur systematics, in 1967 Oskar Kuhn placed Campylognathoides in its own subfamily within the Rhamphorhynchidae, the Campylognathoidinae.[16] However, in 1974 Peter Wellnhofer concluded that it was placed in a more basal position in the phylogenetic tree, below the Rhamphorhynchidae.[17] In the early twenty-first century this was confirmed by the first extensive exact cladistic analyses. In 2003 both David Unwin and Alexander Kellner introduced a clade Campylognathoididae; within Unwin's terminology this clade is the sister clade of the Breviquartossa within the Lonchognatha;[18] applying Kellner's terminology it is the most basal offshoot within the Novialoidea.[19] There is no material difference between the two positions.

According to the analyses Campylognathoides would be closely related to Eudimorphodon, to which it is similar in skull, sternum and humerus form. This was confirmed by Padian in 2009, though Padian also pointed out several basal features present in Eudimorphodon but lacking in Campylognathoides.[20] In 2010 an analysis was published by Brian Andres showing that Eudimorphodon together with Austriadactylus formed a very basal clade, leaving Campylognathoides as the only known member of the Campylognathoididae.[21] Other recent phylogenetic analyses confirm these results, and suggest that Campylognathoides is more derived than all Triassic-aged pterosaurs, as well as the Early Jurassic Dimorphodon and Parapsicephalus.[22][23] Campylognathoides is the basalmost member of the Novialoidea which is defined as a node-based taxon consisting of the last common ancestor of Campylognathoides, Quetzalcoatlus and all its descendants.[24]


Traditionally a piscivorous lifestyle is attributed to Campylognathoides, as to most pterosaurs; in this case supported by the provenance of the finds from marine sediments and the very long wings. Pterosaurologist Kevin Padian, however, has suggested that, in view of the stout short teeth, ideal for delivering a piercing bite, the form might have been a predator of small terrestrial animals instead.[25] Conversely, Mark Witton suggests the construction of Campylognathoides' extremely robust forelimbs, with proportionally long wing fingers, could be a specialization for a fast aerial lifestyle comparable to those of falcons and mastiff bats.[26]

See also


  1. ^ Barrett, P. M., Butler, R. J., Edwards, N. P., & Milner, A. R. (2008). "Pterosaur distribution in time and space: an atlas". Zitteliana, 61-107. [1]
  2. ^ a b c d e "Campylognathoides." In: Cranfield, Ingrid (ed.). The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures. London: Salamander Books, Ltd. Pp. 292-295.
  3. ^ Padian (2009), pp. 100-101
  4. ^ Wellnhofer, P. & Vahldiek, B.-W. (1986). "Ein Flugsaurier-Rest aus dem Posidonienschiefer (Unter-Toarcium) von Schandelah bei Braunschweig", Paläontologische Zeitschrift, 60: 329-340
  5. ^ Padian (2009), p. 103
  6. ^ Quenstedt, F. A. (1858). "Ueber Pterodactylus liasicus", Jahrbuch des Vereins vaterländischer Naturkundler in Württemberg, 14:299-336
  7. ^ Plieninger, F. (1894). "Campylognathus Zittelli. Ein neuer Flugsaurier aus dem Oberen Lias Schwabens", Palaeontographica, 41: 193-222
  8. ^ Plieninger, F. (1901). "Beiträge zur Kenntnis der Flugsaurier", Palaeontographica, 48: 65–90
  9. ^ Plieninger, F. (1906). "Notizen über Flugsaurier aus dem Lias Schwabens", Centralblatt für Mineralogie, 10: 290–293
  10. ^ Plieninger, F. (1907) "Die Pterosaurier der Juraformation Schwabens", Palaeontographica, 53: 209-313
  11. ^ Strand, E. (1928). "Miscellanea nomenclatorica Zoologica et Palaeontologica", Archiv für Naturgeschichte, 92: 30-75
  12. ^ Jain, S. L. (1974) "Jurassic Pterosaur from India", Journal of the Geological Society of India, 15 (3): 330-335
  13. ^ Padian (2009), p. 98
  14. ^ Padian (2009), p. 71, 105
  15. ^ Nopcsa, F. v. (1928). "The genera of reptiles". Palaeobiologica, 1: 163-188
  16. ^ Kuhn, O. (1967). Die fossile Wirbeltierklasse Pterosauria. Krailling: Oeben-Verlag. 52 pp
  17. ^ Wellnhofer, P. (1974). "Campylognathoides liasicus (Quenstedt), an Upper Liassic pterosaur from Holzmaden — The Pittsburgh specimen", Annals of Carnegie Museum, 45: 5-34
  18. ^ Unwin, D. M. (2003). "On the phylogeny and evolutionary history of pterosaurs". Pp. 139-190 in: Buffetaut, E. and Mazin, J.-M., eds. Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217. Geological Society of London
  19. ^ Kellner, A. W. A. (2003). "Pterosaur phylogeny and comments on the evolutionary history of the group". Pp. 105-137 in: Buffetaut, E. and Mazin, J.-M., eds. Evolution and Palaeobiology of Pterosaurs. Geological Society Special Publications 217. Geological Society of London
  20. ^ Padian (2009), pp. 103-104
  21. ^ Brian Andres; James M. Clark & Xu Xing. (2010). "A new rhamphorhynchid pterosaur from the Upper Jurassic of Xinjiang, China, and the phylogenetic relationships of basal pterosaurs", Journal of Vertebrate Paleontology, 30(1): 163-187
  22. ^ Wang X.; Kellner, A. W. A.; Jiang S.; Meng X. (2009). "An unusual long-tailed pterosaur with elongated neck from western Liaoning of China". Anais da Academia Brasileira de Ciências. 81 (4): 793–812. doi:10.1590/S0001-37652009000400016.
  23. ^ Andres, Brian Blake (2010). Systematics of the Pterosauria. Yale University. p. 366. A preview that shows the cladogram without clade names
  24. ^ Kellner, A. W. A., (2003): Pterosaur phylogeny and comments on the evolutionary history of the group. pp. 105-137. — in Buffetaut, E. & Mazin, J.-M., (eds.): Evolution and Palaeobiology of Pterosaurs. Geological Society of London, Special Publications 217, London, 1-347
  25. ^ Padian (2009), p. 105
  26. ^ Witton, Mark P. (2013), Pterosaurs: Natural History, Evolution, Anatomy


  • Dinosaurs and other Prehistoric Creatures, edited by Ingrid Cranfield, 2000 Salamander Books Ltd pg 285-286.
  • Quenstedt, F. A. 1858 "Ueber Pterodactylus liasicus", Jahreshefte des Vereins für vaterländische Naturkunde in Württemberg 14, 299–310 & pl. 2.
  • Plieninger, F. 1907 "Die Pterosaurier der Juraformation Schwabens", Paläontographica 53, 209–313 & pls 14–19.
  • Wellnhofer, P. 1974 "Campylognathoides liasicus (Quenstedt), an Upper Liassic pterosaur from Holzmaden - The Pittsburgh specimen", Ann. Carnegpterus. 45 (2), 5–34.
  • Plieninger, F. 1894 "Campylognathus Zitteli, ein neuer Flugsaurier aus dem obersten Lias Schwabens", Paläontographica 41, 193–222 & pl. 19.
  • Jain, 1974, "Jurassic Pterosaur from India", Journal of the Geological Society of India, vol.15, Pt.3 pp. 330–335
  • Padian, K. 2009. The Early Jurassic Pterosaur Dorygnathus banthenis (Theodori, 1830) and The Early Jurassic Pterosaur Campylognathoides Strand, 1928, Special Papers in Palaeontology 80, Blackwell ISBN 978-1-4051-9224-8
1928 in paleontology

Paleontology or palaeontology is the study of prehistoric life forms on Earth through the examination of plant and animal fossils. This includes the study of body fossils, tracks (ichnites), burrows, cast-off parts, fossilised feces (coprolites), palynomorphs and chemical residues. Because humans have encountered fossils for millennia, paleontology has a long history both before and after becoming formalized as a science. This article records significant discoveries and events related to paleontology that occurred or were published in the year 1928.


Austriadactylus is a genus of "rhamphorhynchoid" pterosaur. The fossil remains were unearthed in Late Triassic (middle Norian age) rocks of Austria.

The genus was named in 2002 by Fabio Marco Dalla Vecchia e.a.. The type species is Austriadactylus cristatus. The genus name is derived from Latin Austria and Greek daktylos, "finger", in reference to the wing finger of pterosaurs. The specific epithet means "crested" in Latin, a reference to the skull crest.

The genus is based on holotype SMNS 56342, a crushed partial skeleton on a slab, found in an abandoned mine near Ankerschlag in Tyrol, in the Norian Seefelder Beds. The counterslab has been lost and with it some of the bone. The fossil consists of the skull, lower jaws, some vertebrae, parts of the limbs and pelvic girdle, and the first part of the tail.

The elongated skull has a length of 11 cm. It carried a bony crest that widened as it descended towards the snout, up to height of 2 cm. The triangular nares formed the largest skull openings. The also triangular fenestrae antorbitales are smaller than the orbits. The teeth differ in shape and the species was thus heterodont. Most teeth are small and tricuspid or three-pointed. In the front of the upper jaw five larger recurved teeth with a single point form a prey grab; six or seven such teeth are also interspersed with the smaller teeth more to the back of the mouth. There are at least seventeen and perhaps as much as 25 tricuspid teeth in the upper jaw, for a total of perhaps 74 teeth of all sizes in the skull. The number of teeth in the lower jaws cannot be determined.

The flexible tail did not have the stiffening rod-like vertebral extensions present in other basal pterosaurs. The wingspan has been estimated at about 120 cm.

Austriadactylus was in 2002 assigned by the describers to a general Pterosauria incertae sedis, but some later analyses showed it to have been related to Campylognathoides and Eudimorphodon in the Campylognathoididae. It has even been suggested it was a junior synonym of Eudimorphodon, though perhaps a distinct species in that genus. The following phylogenetic analysis follows the topology of Upchurch et al. (2015).


Averostra, or "bird snouts", is a clade that includes most theropod dinosaurs that have a promaxillary fenestra (fenestra promaxillaris), an extra opening in the front outer side of the maxilla, the bone that makes up the upper jaw. Two groups of averostrans, the Ceratosauria and the Orionides, survived into the Cretaceous period. When the Cretaceous–Paleogene extinction event occurred, ceratosaurians and two groups of orionideans within the clade Coelurosauria, the Tyrannosauroidea and Maniraptoriformes, were still extant. Only one subgroup of maniraptoriformes, Aves, survived the extinction event and persisted to the present day.


Avetheropoda, or "bird theropods", is a clade that includes carnosaurians and coelurosaurs to the exclusion of other dinosaurs.


Caelestiventus ( sə-LES-tih-VEN-təs, meaning "heavenly wind") is a pterosaur genus from the Late Triassic (Norian or Rhaetian) found in western North America. The type species, Caelestiventus hanseni, honors Robin Hansen, the Bureau of Land Management geologist (BLM), who facilitated access to the excavation site.

Caelestiventus is important because it is the sole example of a desert-dwelling non-pterodactyloid pterosaur and is 65 million years older than other known desert-dwelling pterosaurs. Additionally, it shows that even the earliest pterosaurs were morphologically and ecologically diverse and that the Dimorphodontidae originated in the Triassic Period.


Cerapoda ("ceratopsians and ornithopods") is a clade of the dinosaur order Ornithischia.


Dinosauriformes is a clade of archosaurian reptiles that include the dinosaurs and their most immediate relatives. All dinosauriformes are distinguished by several features, such as shortened forelimbs and a partially to fully perforated acetabulum, the hole in the hip socket traditionally used to define dinosaurs. The oldest known member is Asilisaurus, dating to about 245 million years ago in the Anisian age of the middle Triassic period.


Dorygnathus ("spear jaw") was a genus of pterosaur that lived in Europe during the Early Jurassic period, 180 million years ago when shallow seas flooded much of the continent. It had a short 1.5 meters (4.9 feet) wingspan, and a relatively small triangular sternum, which is where its flight muscles attached. Its skull was long and its eye sockets were the largest opening therein. Large curved fangs that "intermeshed" when the jaws closed featured prominently at the front of the snout while smaller, straighter teeth lined the back. Having variable teeth, a condition called heterodonty, is rare in modern reptiles but more common in primitive pterosaurs. The heterodont dentition in Dorygnathus is consistent with a piscivorous (fish-eating) diet. The fifth digit on the hindlimbs of Dorygnathus was unusually long and oriented to the side. Its function is not certain, but the toe may have supported a membrane like those supported by its wing-fingers and pteroids. Dorygnathus was according to David Unwin related to the Late Jurassic pterosaur, Rhamphorhynchus and was a contemporary of Campylognathoides in Holzmaden and Ohmden.

Graphical timeline of pterosaurs

Timeline showing the development of the extinct reptilian order Pterosauria from its appearance in the late Triassic period to its demise at the end of the Cretaceous, together with an alphabetical listing of pterosaur species and their geological ages.


Jeholosaurids were herbivorous neornithischian dinosaurs from the Cretaceous Period (Aptian - Santonian, with a possible Campanian record) of Asia. The family was first proposed by Han et al. in 2012. The jeholosaurids were defined as those ornithischians more closely related to Jeholosaurus shangyuanensis than to Hypsilophodon foxii, Iguanodon bernissartensis, Protoceratops andrewsi, Pachycephalosaurus wyomingensis, or Thescelosaurus neglectus. The Jeholosauridae includes the type genus Jeholosaurus and Yueosaurus.


Jingshanosaurus (meaning "Jingshan lizard") is a genus of sauropodomorph dinosaurs from the early Jurassic period.

Kota Formation

The Kota Formation is a geological formation in India. The dates for the Kora Formation are uncertain, but it dates from the Early Jurassic to Early Cretaceous, and is split into a Lower Member and Upper Member. The lower members is though to be Hettangian-Pliensbachian.


The Melanorosauridae were a family of sauropodomorph dinosaurs which lived during the Late Triassic and Early Jurassic. The name Melanorosauridae was first coined by Friedrich von Huene in 1929. Huene assigned several families of dinosaurs to the infraorder "Prosauropoda": the Anchisauridae, the Plateosauridae, the Thecodontosauridae, and the Melanorosauridae. Since then, these families have undergone numerous revisions. Galton and Upchurch (2004) considered Camelotia, Lessemsaurus, and Melanorosaurus members of the family Melanorosauridae. A more recent study by Yates (2007) indicates that the melanorosaurids were instead early sauropods.


Neotheropoda (meaning "new theropods") is a clade that includes coelophysoids and more advanced theropod dinosaurs, and the only group of theropods who survived the Triassic–Jurassic extinction event. Yet all of the neotheropods became extinct during the early Jurassic period except for Averostra.


Novialoidea (meaning "new wings") is an extinct clade of macronychopteran pterosaurs that lived from the latest Early Jurassic to the latest Late Cretaceous (early Toarcian to late Maastrichtian age), their fossils having been found on all continents except Antarctica. It was named by Alexander Wilhelm Armin Kellner in 2003 as a node-based taxon consisting of the last common ancestor of Campylognathoides, Quetzalcoatlus and all its descendants. This name was derived from Latin novus "new", and ala, "wing", in reference to the wing synapomorphies that the members of the clade possess. Unwin (2003) named Lonchognatha in the same issue of the journal that published Novialoidea (Geological Society of London, Special Publications 217) and defined it as Eudimorphodon ranzii, Rhamphorhynchus muensteri, their most recent common ancestor and all its descendants (as a node-based taxon). Under Unwin's and Kellner's phylogenetic analyses (where Eudimorphodon and Campylognathoides form a family that basal to both Rhamphorhynchus and Quetzalcoatlus), and because Novialoidea was named first (in pages 105-137, while Lonchognatha was named in pages 139-190), Lonchognatha is an objective junior synonym of the former. However, other analyses find Lonchognatha to be valid (Andres et al., 2010) or synonymous with the Pterosauria (Andres, 2010 and Andres, in press).


Orionides is a clade of tetanuran theropod dinosaurs from the Middle Jurassic to the Present. The clade includes most theropod dinosaurs, including birds.


Orodrominae is a subfamily of parksosaurid dinosaurs from the Cretaceous of North America and Asia.


Riojasauridae is a family of sauropod-like dinosaurs from the Upper Triassic. It is known primarily from the genera Riojasaurus and Eucnemesaurus. Sites containing Riojasauridae include the Lower Elliot Formation of Orange Free State, South Africa (where fossils of Eucnemesaurus have been found), and Ischigualasto, in La Rioja Province, Argentina ( where fossils of Riojasaurus have been recovered).


Xixiposaurus is a genus of prosauropod dinosaur which existed in what is now Lower Lufeng Formation, China during the lower Jurassic period. It was first named by Sekiya Toru in 2010 and the type species is Xixiposaurus suni.


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