Ornithocheiromorphs Temporal range:Valanginian-Turonian ~140–92.5 Ma PreꞒ Ꞓ O S D C P T J K Pg N
Reconstructed skeleton ofTropeognathus in theNational Museum of Brazil
Skeletal cast ofMaaradactylus spielbergi in theNaturalis Biodiversity Center
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Order:	†Pterosauria
Suborder:	†Pterodactyloidea
Clade:	†Pteranodontoidea
Clade:	†Ornithocheiromorpha Andreset al., 2014
Subgroups
†Serradraco †Unwindia †Boreopteridae(relationships uncertain) †Lonchodectidae(relationships uncertain) †LanceodontiaAndreset al., 2014 †Draigwenia †IstiodactyliformesKellneret al., 2019 †Hongshanopterus †Linlongopterus †Yixianopterus †Istiodactylidae †Mimodactylidae †OrnithocheiriformesKellneret al., 2019 †Brasileodactylus †Ornithocheirus(Ornithocheiridae?) †Anhangueridae †Hamipteridae †Targaryendraconia

Ornithocheiromorpha (fromAncient Greek, meaning "bird hand form") is a group ofpterosaurs within thesuborderPterodactyloidea.Fossil remains of this group date back from theEarly toLate Cretaceousperiods (Valanginian toTuronian stages), around 140 to 92.5 million years ago. Ornithocheiromorphs have been discovered worldwide exceptAntarctica, though mostgenera have been recovered inEurope,Asia andSouth America.^[1] They were the most diverse and successful pterosaurs during the Early Cretaceous, but throughout the Late Cretaceous they were replaced bypteranodontians andazhdarchoids. The Ornithocheiromorpha was defined in 2014 by Andres and colleagues, and they made Ornithocheiromorpha the most inclusiveclade containingOrnithocheirus, but notPteranodon.^[2]

Ornithocheiromorphs are considered to be some of the largest animals to have ever flown. Members of this group are also regarded to have some of the largest pterosaurwingspans, such as the one estimated for the hugeTropeognathus, though still not as large as those estimated for theazhdarchids, which may have reached up to 12 meters (39 ft).^[3] When ornithocheiromorphs first appeared, they were initiallyscavengers, consisting in a more terrestrial setting, but their success had made them the top predators of the skies, as well as the most common type offish-eating pterosaur throughout the early Late Cretaceous. Somepaleontologists also consider ornithocheiromorphs an earlier step of evolution to thepteranodontians, this is due to the similar flying techniques and flight locomotions, as well as their diet, which mainly consisted of fish, and therefore also hunted very similarly. Ornithocheiromorphs also flew like soaring birds, keeping their wings stretched and rarely flapping.

The first specimens of ornithocheiromorphs were unearthed at a chalk pit nearBurham inKent,England. In 1846,BritishpaleontologistJames Scott Bowerbank named and described the remains found asPterodactylus giganteus, as it was common at that time to assign any new described pterosaur species toPterodactylus.^[4] In the same chalk pit asP. giganteus, two other pterosaur species were discovered. The first was named in 1851 by Bowerbank asPterodactylus cuvieri,^[5] in honor of the prominent Germannaturalist andzoologistGeorges Cuvier, while the second was described in the same year by British paleontologist SirRichard Owen asPterodactylus compressirostris.^[6]P. compressirostris later became the type species of a newly created genus calledLonchodectes (meaning "lance biter") in a review by English paleontologistReginald Walter Hooley in 1914.^[7] Confusingly, this species was also long regarded, incorrectly, as the type species ofOrnithocheirus.^[8]

In 1861, further pterosaur specimens were found in theUK, and were given the new speciesPterodactylus simus by Owen.^[9] British paleontologistHarry Govier Seeley then created the new genusOrnithocheirus for the new species in the same year, the generic name translating as "bird hand" is due to the notion of the time that pterosaurs were the ancestors of modern birds. In 1870, Seeley reassigned the speciesPterodactylus cuvieri asOrnithocheirus cuvieri.^[10][7] In 1874, Richard Owen proposed two new genera,Coloborhynchus, meaning "maimed beak", andCriorhynchus, meaning "ram beak". WhileColoborhynchus consisted in a totally new type species,C. clavirostris, as well as two other species reassigned fromOrnithocheirus,Criorhynchus consisted entirely of formerOrnithocheirus species, includingO. simus, which was later reassigned by Owen asCriorhynchus simus.^[11]

In 2013, Brazilian paleontologistsTaissa Rodrigues &Alexander Kellner made a deeper analysis on the speciesPterodactylus cuvieri. In the analysis, they stated that it needed a separate genus, and assigning it toOrnithocheirus was inappropriate, therefore, they created the new genus calledCimoliopterus, with the new resulting combinationCimoliopterus cuvieri. In the same study, Rodrigues & Kellner also reviewed the speciesPterodactylus giganteus, and reassigned it to a newly created genus calledLonchodraco, this resulted in a new combination calledLonchodraco giganteus.^[12]

In 1887, Seeley had described new fossil remains from theIsle of Wight, an island off the coast of southern England. He thought it belonged to some kind of bird-like creature, which he named itOrnithodesmus cluniculus.^[13] Seeley also reported another specimen found on the same site. He then considered it another species ofOrnithodesmus. In 1901, Seeley named this new species asO. latidens, meaning "wide tooth".^[14] Later, Reginald Hooley discussedO. latidens in detail, based on specimens he had found, which ledOrnithodesmus to be placed within a new family called Ornithodesmidae.^[15] PaleontologistCharles William Andrews however, had expressed doubts as to whetherO. latidens belonged in the genusOrnithodesmus, as the vertebrae of the specimen of that genus was based on differed markedly from those of Hooley's specimen.^[16]

In 1993, the British paleontologistsStafford C. Howse andAndrew C. Milner concluded that the holotype sacrum and only specimen ofO. cluniculus didn't belong to a pterosaur, but instead to amaniraptorantheropod dinosaur. They also pointed out that no detailed attempts had been made to compare the sacrum ofO. cluniculus with those of pterosaurs, and thatO. latidens had in effect been treated as the type species of the genusOrnithodesmus.^[17] Howse, Milner, and David Martill in 2001, moved "O."latidens to a new genus calledIstiodactylus. They had also named a new family calledIstiodactylidae, withIstiodactylus as the only member.^[18]

Other important ornithocheiromorph discoveries include the anhangueridsTropeognathus andAnhanguera from theRomualdo Formation inBrazil.^[19][20]Tropeognathus was described with its type species,T. mesembrinus in 1987 byGerman paleontologistPeter Wellnhofer. The generic name is derived fromGreek τρόπις,tropis, meaning "keel", and γνάθος,gnathos, meaning "jaw". Thespecific name is derived fromKoinemesembrinos, "of the noontide", simplified as "southern", in reference to the provenance from theSouthern hemisphere. The description then led to an enormous taxonomic confusion.^[21] In 1989,Brazilian paleontologist Alexander Kellner considered it anAnhanguera mesembrinus,^[22] then aColoborhynchus mesembrinus by Veldmeijer in 1998,^[23] and then aCriorhynchus mesembrinus in 2001 by German paleontologistMichael Fastnacht.^[24]T. mesembrinus was then considered ajunior synonym ofOrnithocheirus simus by British paleontologistDavid Unwin in 2001, but he then proposed anOrnithocheirus mesembrinus in 2003.^[25][26] In 2013 however, Taissa Rodrigues and Alexander Kellner concluded thatTropeognathus would be valid again, and containing onlyT. mesembrinus, the type species.^[12]

A discovery in Asia, specifically northwesternChina, was reported in 2006. The lake sediments allowed an exceptional preservation of fossils, and therefore paleontologists Qiu Zhanxiang and Wang Banyue started official excavations. Part of the findings consisted of dense concentrations of pterosaur bones, associated with soft tissues and eggs. In 2014, a new species was named and described:Hamipterus tianshanensis. It was named by Wang Xiaolin, Alexander Kellner, Jiang Shunxing, Wang Qiang, Ma Yingxia, Yahefujiang Paidoula, Cheng Xin, Taissa Rodrigues, Meng Xi, Zhang Jialiang, Li Ning, and Zhou Zhonghe. The generic nameHamipterus combines that of theHami region, with the wordpteron, meaning "wing", and the specific name refers to the provenance from theTian Shan, a mountain range.^[27]

Ornithocheiromorphs were large pterosaurs, with wingspans normally ranging between 3 and 6 meters (9.8 and 19.7 ft).^[3]Istiodactylus for example, had awingspan ranging from 4.3 to 5 meters (14 to 16 ft), with the most complete known skull estimated to have been about 45 centimeters (1.48 ft) in length, based on a long-lost fragment of its jaw reported in 2012.^[28] Though its jaws measured only 28.5 centimeters (11.2 in), which was less than 80 percent of the skull's length.^[16] Anhanguerids and were typically larger than others of the group and were more successful within thefood chain rather than other ornithocheiromorphs, one reason is because of their large size, for example,Tropeognathus mesembrinus, had a normal wingspan of about 8.26 meters (27.1 ft), and 8.70 meters (28.5 ft) as the maximum estimate.^[29] Another species which was impressively large isColoborhynchus capito, with a total skull length that could have been up to 75 centimeters (2.46 ft), leading to an estimated wingspan of 7 meters (23 ft).^[30] However, this species may belong to a different genus calledNicorhynchus.^[31]

Most anhanguerids bore distinctive convex "keeled" crests on their snout and underside of theirmandible, this was well developed in several genera such asTropeognathus.^[32] The similarAnhanguera possessed jaws that were tapered in width, but expanded into a broad, spoon-shaped rosette at the tip. The jaws are distinguished from its relatives by several differences in the crest and teeth: unlike its close relativesColoborhynchus andOrnithocheirus, the crest on the upper jaw ofAnhanguera didn't begin at the tip of the snout, therefore, it was set farther back on the skull.^[33]

Other anhanguerids likeCearadactylus had its first preparations with many serious mistakes: the front of the snout and the lower jaws were confused leading to a reconstruction in which the anterior part of the head was upside down.^[34] Some of the teeth were extensively restored and enlarged until the wider front of the jaws showed very large and robust teeth projecting outward. With this arrangement, the maxilla was kinked, and its interlockingteeth suggested thatCearadactylus had apiscivourous diet, allowing the animal to keep hold of slippery fish.^[35] Another smaller genus similar toCearadactylus isGuidraco. Its holotypeskull has a length of 38 centimeters (15 in), which makes it smaller than other genera. The skull is very elongated however, and a hollow profile is seen, but not very pointed, as the upper edge and the line of the jaw run nearly parallel over most of their length.^[36]

Even though most ornithocheiromorphs didn't have acranial crest like the closely related pteranodontids, there were some exceptions, this includedCaulkicephalus andLudodactylus.^[37]Caulkicephalus had a rounded snout, very similar to that ofOrnithocheirus andAnhanguera, and therefore it is placed within either Anhangueridae or Ornithocheiridae, depending on the author.^[38][39]Caulkicephalus was also a large pterosaur, with wingspan estimates of around 5 meters (16 ft).^[39]

The vertebral column of ornithocheiromorphs was heavily pneumatized by an extensive system of air sacs, leaving prominent pneumatic foraminae.^[40] The neck of ornithocheiromorphs was typically relatively long and robust, being longer than the torso in some derived clades.^[41] The neural spines of ornithocheiromorph cervical vertebrae were generally tall and spikelike.^[36][33] In some genera such asTropeognathus andIstiodactylus, up to six dorsal vertebrae are fused into anotarium.^[42] In some genera such asAnhanguera, four to seven sacral vertebrae are fused into asynsacrum.^[33] The tail is not well known in ornithocheiromorphs, however.Zhenyuanopterus, which is known for having 13 caudal vertebrae, formed one of the longest tails of any pterodactyloid.^[43]Anhanguera, another well-known genus, had a shorter tail, with broad caudal vertebrae that bore a "duplex" cross-section similar toPteranodon.^[33]

Thepelvis of ornithocheiromorphs was of moderate size compared to the body as a whole, similar to otherornithocheiroids. The threepelvic bones were often fused, as seen in many species such asAnhanguera santanae, theilium was long and low, and its front and rear blades projected horizontally beyond the edges of the lower pelvic bones.^[44] Despite the structure length, the processes of these rod-like forms indicate that the hindlimb muscles attached to them were limited in strength.^[45] Thepubic bone was fused with the broad ischium into an ischiopubic blade, resulting in a narrow build. Sometimes, the blades of both sides were fused, closing the pelvis from below and forming the pelvic canal. The front of the pubic bones was also articulated with a unique structure, resulting in a pair of prepubic bones within. This formed a cusp covering the rear belly, and was located between the pelvis and the belly ribs. Thehip joint of ornithocheiromorphs was not perforated and allowed considerable mobility to the leg, and suggests that it was vertical, as therefore had a function in breathing, compensating the relative rigidity of the chest cavity.^[46][44]

Several studies show that ornithocheiromorphs were less derived than the toothless pteranodontids such asPteranodon, and based on the different evolutionary changes, they therefore need to be grouped in a differentclade thanPteranodon, though still withinPteranodontoidea.^[47] In 2003, David Unwin considered the family Istiodactylidae to group with the toothless Pteranodontidae, within the groupOrnithocheiroidea,^[26] but Alexander Kellner however, grouped it with the toothed Anhangueridae instead, resulting in a more understandable change of evolution between the two toothed families.^[48]

Brian Andres and colleagues found the families Istiodactylidae, Ornithocheiridae, and Anhangueridae to form a group in 2014, which he calledLanceodontia. The clade includes most families of ornithocheiromorphs, however they excluded the poorly-known familyLonchodectidae from membership in this clade. In their analysis, they also included the familyBoreopteridae within the cladeAnhangueria, though placed in a more basal position, while also containing the genusGuidraco.^[2] In 2018 however, Nicholas Longrich and colleagues found Boreopteridae outside Anhangueria as the sister taxon of the family Lonchodectidae, both groups placed as basal members of the Ornithocheiromorpha.^[49]

Within Lanceodontia, there is a more restrictive clade calledOrnithocheirae, which has a more convoluted taxonomic history. It was named byHarry Seeley in1870 as a family that containsOrnithocheirus and its relatives. The name was emended toOrnithocheiridae, to match the requirements of theICZN Code that a family-ranked clade should end with an "-idae"suffix. Brian Andres (2010) in his review of pterosaurphylogeny, defined the name Ornithocheirae phylogenetically, as anode-based taxon consisting of the last common ancestor ofAnhanguera andOrnithocheirus and all its descendants.^[50] Thus Ornithocheirae is defined to include two families, theAnhangueridae and theOrnithocheiridae, following the opinion ofAlexander Kellner^[51] and Andres that these families should not be synonymized based on their original phylogenetical definitions.^[50] Subsequent studies in 2019 have found Ornithocheirae to be a more inclusive group containing both Anhangueria (see below) andTargaryendraconia.

Another constituent clade of Ornithocheiromorpha isAnhangueria. Anhangueria was named by paleontologists Taissa Rodrigues andAlexander Kellner in a review ofOrnithocheirus species in 2013, they defined the clade as abranch-based taxon consisting of all pteranodontoids more closely related toAnhanguera blittersdorffi than toIstiodactylus latidens andCimoliopterus cuvieri.^[12] Anhangueria originally only contained thegeneraBrasileodactylus,Camposipterus,Cearadactylus,Ludodactylus as well as the familyAnhangueridae,^[12] however, recent analyses had recovered the familyHamipteridae within this clade as well.^{[52][53][38][31]} In 2014, paleontologist Brian Andres and colleagues assigned more groups and genera within this clade, this includedGuidraco, the subfamilyBoreopterinae, and the cladeOrnithocheirae, which was further divided into the families Anhangueridae andOrnithocheiridae.^[2] This topology was later followed by a few other studies.^[49][54]

In 2019, Alexander Kellner and colleagues described the new ornithocheiromorphMimodactylus, and used their analysis to name a new clade,Istiodactyliformes. They defined this clade as the most inclusive clade containingIstiodactylus but notAnhanguera. Under their analysis, this new clade included the familyIstiodactylidae,Mimodactylidae, and the genusHongshanopterus.^[55] Andres and colleagues later named a cladeOrnithocheiriformes and defined it in opposition to Istiodactyliformes. Its given definition was the most inclusive clade containingAnhanguera, but notIstiodactylus.^[56]

Some of these clades have been recovered by various authors since their definition. A table summarizing all of these clades is shown below.

There are competing theories of ornithocheiromorphphylogeny (evolutionary relationships). Below is acladogram showing the results of aphylogenetic analysis presented by Longrichet al. (2018). In the analysis, they placed the genusHongshanopterus as a basal member.^[49]

In 2019, several new species of ornithocheiromorphs were found, and the former speciesOrnithocheirus wiedenrothi was renamed asTargaryendraco wiedenrothi.^[38] The description ofIberodactylus inSpain also made some paleontologists reclassify the genusHamipterus in a newly named family calledHamipteridae.^[52] The ornithocheiridCimoliopterus was also reclassified as well, and it is currently grouped withAetodactylus andCamposipterus in the cladeTargaryendraconia, specifically to its own family, theCimoliopteridae.^[38] However, an analysis by Jacobset al. (2019) recovers bothCamposipterus andCimoliopterus within the Ornithocheiridae again, using a new data matrix not includingTargaryendraco.^[58] The previously recovered basaleupterodactyloidHaopterus was reclassified due to the description ofMimodactylus, and is placed in a new family calledMimodactylidae.^[53] However, a more recent analysis using the data in the description ofMimodactylus has foundHaopterus as a basal member of the more inclusive group Istiodactyliformes.^[59] Many recent analyses have also recovered several ornithocheirids, includingTropeognathus,Coloborhynchus, andCaulkicephalus within the family Anhangueridae, meaning that they were more closely related toAnhanguera than toOrnithocheirus.^{[52][38][59][53][31]}

Later, in 2019, a different phylogenetic analysis, this time conducted by Borja Holgado and colleagues, focused on derivedpterodactyloids with particular emphasis on anhanguerians. In this analysis, Anhangueria consisted ofCamposipterus and the families Anhangueridae and Hamipteridae, the clade Ornithocheirae was recovered as a more inclusive group consisting ofOrnithocheirus andCimoliopterus asbasal members, as well as the Anhangueria in the most derived position. This classification is supported by the expansion of thepremaxillary tip (a spoon-like expansion at the end of the snout) with a high jaw end.^[52] Many analyses afterwards have followed this concept,^[53][60] although some had recoveredCamposipterus outside the Anhangueria (within the cladeTargaryendraconia), but still within the Ornithocheirae.^[38][31]

Below iscladogram following a topology recovered by Brian Andres, using the most recent iteration of his data set (Andres, 2021).^[56]

In their description ofLingyuanopterus in 2022, Xu Yizhi and colleagues recovered the following phylogeny based on the earlier work of Borja Holgado and Rodrigo Pêgas.

Ornithocheiromorphs were originally regarded aspiscivorous creatures, feeding mainly on small and mid-sized fish.^[35] Some paleontologists even suggested details on how these pterosaurs caught fish, some of which included dipping theirbeaks close to the water for prey.^[35] Hooley for example, found that the beak of the well knownIstiodactylus was similar to those of birds such asherons,storks, andskimmers, and suggested thatIstiodactylus probably fed on fish, this was mainly based on his 1913 jaw reconstruction of the animal.^[15] In 1991, Peter Wellnhofer compared the jaw endings ofIstiodactylus with those of aduck, but he then noticed that it wasn't a "duck-billed pterosaur" or anything similar, even though it was popularly called that way.^[61] An analysis by Witton in 2012 found that the teeth ofIstiodactylus were unlike the recurved and enlarged teeth seen in the more derived ornithocheirids such asOrnithocheirus, he instead pointed out that it was more "razor-edged" and better suited for carrion rather than fish.^[28]

Another ornithocheiromorph that possessed similar features toIstiodactylus isLiaoxipterus, which is known from a skull with several unique traits, including numerous peg-like teeth. The shape of its teeth indicated thatLiaoxipterus was a possibleinsectivore,^[62] though this conclusion isn't considered entirely accurate.^[63]

Anhanguerids likeTropeognathus andColoborhynchus are considered to be fish-eaters, and had longer and sharper teeth compared to the more rounded teeth ofIstiodactylus, though this is still sometimes disputed.^[35] Another difference that can be seen in more primitive ornithocheiromorphs their eyes being proportionally smaller compared to the assumed predatory and more advanced anhanguerids, this again adds to the fact that the moreprimitive groups were most likely scavengers, and later got more successful within the food chain, leading the later, more advanced groups to be thedominant fish hunters during the early Late Cretaceous.^[64]

Ornithocheiromorphs, like other pterosaurs, are considered to have been skilled fliers as well as swift at moving on the ground. Footprints from several species show that most pterosaurs did not sprawl their limbs to a large degree, as in modern reptiles, but rather held the limbs relatively erect when walking, likedinosaurs. While footprints are yet to be known, it is likely that ornithocheiromorphs also walked erect.^[65] Compared to other earlier pterosaurs such as rhamphorhynchids, ornithocheiromorphs had unusually unevenlimb proportions, with the forelimbs resulting in a much longer scale compared to the hind limbs. Their close relatives, the pteranodontids, were also found with similar features, though they more likely flew like modern-dayalbatrosses rather than anything else. Paleontologists also suggest that they most likely spend long stretches of time sea fishing, traveling very long distances without flapping while at the same time flying close the surface of the water with exploited wind speed, and without the necessity ofthermals.^[66] This would likely have required them to use unique modes of locomotion compared to other pterosaurs, this can already be seen in earlier evolutions such asIstiodactylus andNurhachius, with powerful musculature attachments and well-developed pectoral and upper arm bones.^[28][67] It is also possible that ornithocheiromorphs ran (but not walked) bipedally, or that they used a hopping gait.^[65] Many pterosaur researchers like Mike Habib have noted that the limb proportions of some ornithocheiromorphs such asAnhanguera are consistent with hopping, though the scavenging istiodactylids are probably still the best examples of pterosaurs with a more terrestrial setting.^[68][15]

Even though the ornithocheiromorphs were discovered worldwide, most of them were concentrated in specific places. One of which is the fossil site called Romualdo Formation, which contains an impressive amount of pterosaur fossils. It is a diverseLagerstätte in the larger geologic group called theSantana Group (sometimes called the Santana Formation), which is located in theAraripe Basin of northeastern Brazil, and dates back around 111 and 108 million years ago, during theAlbian stage of the Early Cretaceous. The formation includes many species ofAnhanguera,^[33] several fossil remains of basal ornithocheiromorphs, includingBrasileodactylus,Cearadactylus andUnwindia, as well as the anhangueridsTropeognathus,^[19]Coloborhynchus,Maaradactylus, andAraripesaurus.^[24] These pterosaur genera were just some of the many recovered from the site, which also include thethalassodrominesTupuxuara andThalassodromeus,^[69] as well as thetapejaridTapejara.^[70]

Some other creatures from the formation include the theropodsIrritator,Mirischia andSantanaraptor, and thecrocodylomorphAraripesuchus. The formation also includes several turtle remains, with some specimens referring toSantanachelys,Cearachelys andAraripemys.^[71] A few fish remains were also found within the Romualdo Formation, some of which were referred toBrannerion,Rhinobatos,Rhacolepis,Tharrhias andTribodus. The Santana Group also consists of another Lagerstätte called theCrato Formation, which is not as diverse as the Romualdo Formation, but its fossil remains are still considered important.^[72] This fossil site underlies the Romualdo Formation, and dates back around 115 and 113 million years ago during theAptian stage, meaning that its fossil content is of older age. Similarly, the Crato Formation also contained several species of pterosaurs, including the basal lanceodontiansLudodactylus^[37] andBrasileodactylus, as well as the ornithocheiridArthurdactylus. Other pterosaur genera include the tapejaridsTupandactylus andAymberedactylus,^[70] as well as thechaoyangopteridLacusovagus. The formation also contains other creatures such as theenantiornithineCratoavis, theneosuchianSusisuchus, and several species of fish, includingBelonostomus,Calamopleurus,^[73]Cladocyclus,^[74]Dastilbe^[75] andLepidotes.^[76] These fish genera were suggested to be prey for the pterosaurs that lived in the formation, but fossil remains are limited, so the subject is still controversial.^[35]

Another important fossil site is theWessex Formation in the Isle of Wight, near the coast of England, which dates back around 140 and 125 million years ago (Berriasian toBarremian stages). The formation doesn't contain many fossil remains of pterosaurs compared to the Romualdo Formation, but it is still a very important site. Fossil remains of the istiodactylidIstiodactylus,^[28] the anhangueridCaulkicephalus,^[39] and the tapejaridWightia were found.^[77] The formation is also known for several theropods, including the spinosauridBaryonyx, thetyrannosauroidEotyrannus, thedromaeosauridOrnithodesmus, thecompsognathidAristosuchus as well as theallosauroidNeovenator.^[78][79] Different types of herbivorous dinosaurs likeIguanodon,Polacanthus,Ornithopsis,Mantellisaurus andHypsilophodon were also found within the fossil site.^[80][81] Some other animals from the formation include the neosuchianBernissartia, sea turtles such asHelochelydra andBrodiechelys, the cartilaginous fishHybodus,ray-finned fishes such asBelonostomus,Caturus,Lepidotes andScheenstia, as well as themammalsEobaatar,Loxaulax andYaverlestes.^[76][82][83]

A few fossils reported from theToolebuc Formation andWinton Formation are believed to be from some of the last ornithocheiromorphs, due to the age of the fossil remains, which dated back to the Albian andCenomanian stages of the Cretaceous, and some are even believed to belong to the Turonian stage. The Toolebuc Formation includes several remains of ornithocheiromorphs which are now referred to the generaAussiedraco andMythunga.^[84][85] The formation also includes several herbivorous dinosaurs such as theornithopodMuttaburrasaurus and theankylosaurKunbarrasaurus.^[86][87] Fossil remains of marine animals were also uncovered within the fossil site, and some specimens of which belong to theichthyosaurPlatypterygius, thepliosauridKronosaurus and theelasmosauridEromangasaurus.^[88] Turtle remains from turtles that were proposed to be prey for pterosaurs were also found within the Toolebuc Formation, this included the generaBouliachelys,Cratochelone andNotochelone.^[35] The Winton Formation consisted on a more terrestrial environment, containing severalsauropod dinosaurs likeAustrosaurus,Diamantinasaurus,Savannasaurus andWintonotitan as well as large carnivorous dinosaurs such asAustralovenator^[89] and crocodylomorphs likeIsisfordia. The formation's only pterosaur is the derived genusFerrodraco,^[54] which is also considered as one of the last ornithocheiromorphs, and a close relative ofMythunga.^[85]

• Pterosaur size

• Wellnhofer, Peter (1991).The Illustrated Encyclopedia of Pterosaurs: An illustrated natural history of the flying reptiles of the Mesozoic Era. Crescent Books.ISBN 0-517-03701-7.
• Witton, Mark (2013).Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press.ISBN 978-0-691-15061-1.

• Data related toOrnithocheiromorpha at Wikispecies

• Valanginian first appearances
• Turonian extinctions
• Fossil taxa described in 2014
• Pteranodontoids

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