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tation 18 and the adaptation of this for the public in the illustrated encyclopedia of dinosaurs 19 formerly assigned species and material edit in 1984 argentine paleontologist josé bonaparte and colleagues named kritosaurus australis for hadrosaur bones from the late campanian early maastrichtian los alamitos formation of rio negro patagonia argentina 20 in 2010 this species was thought to be a synonym of secernosaurus koerneri 21 further analysis proved the bones to belong to a new genus thus huallasaurus was named by rozadilla et al 2022 22 in 1990 jack horner and david b weishampel once again separated gryposaurus citing the uncertainty associated with the latter s partial skull horner in 1992 described two more skulls from new mexico that he claimed belonged to kritosaurus and showed that it was quite different from gryposaurus 23 but the following year adrian hunt and spencer g lucas put each skull in its own genus creating anasazisaurus and naashoibitosaurus 24 adrian hunt and spencer g lucas american paleontologists named anasazisaurus horneri in 1993 the name was derived from the anasazi an ancient native american people and the greek word sauros lizard the anasazi were famous for their cliff dwellings such as those in chaco canyon near the location of fossil anasazisaurus remains the term anasazi itself is actually a navajo language word anaasází enemy ancestors the species was named in honor of jack horner the american paleontologist who first described the skull in 1992 the holotype skull and only known specimen was collected in the late 1970s by a brigham young university field party working in san juan county and is housed at byu as byu 12950 24 in 2002 specimen txvp 41917 1 a partial left maxilla recovered from bruja canyon in big bend national park was assigned to kritosaurus as cf k navajovius twenty years later it was discovered to contain a number of useful diagnostic traits that allow it to be described as a new taxon and thus malefica was named by prieto márquez and wagner 2023 25 unnamed specimen from the sabinas basin in mexico assigned to kritosaurus sp by kirkland et al 2006 26 but considered an indeterminate saurolophine by prieto márquez 2013 27 the specimen is informally known as sabinosaurus 28 horner originally assigned the anasazisaurus skull to kritosaurus navajovius 23 but hunt and lucas could not find any diagnostic features in the limited material of kritosaurus and judged the genus to be a nomen dubium since the anasazisaurus skull did have diagnostic features of its own and did not appear to them to share any unique features with kritosaurus it was given the new name anasazisaurus horneri 24 an opinion which was supported by some later authors 13 not all authors have agreed with this thomas e williamson in particular defending horner s original interpretation 4 and several subsequent studies recognized both distinct genera 26 29 a comprehensive study of known kritosaurus material published by albert prieto márquez in 2013 upheld the status of naashoibitosaurus as a distinct genus but found that the type specimens of kritosaurus and anasazisaurus were indistinguishable when comparing overlapping elements i e only those bones preserved in both specimens prieto márquez therefore regarded anasazisaurus as a synonym of kritosaurus but retained it as the distinct species k horneri 27 a partial skeleton from the sabinas basin in mexico was described as kritosaurus sp by jim kirkland and colleagues 26 but considered an indeterminate saurolophine by prieto márquez 2013 27 this skeleton is about 20 larger than other known specimens around 11 meters 36 ft long and with a distinctively curved ischium and represents one of the largest known well documented north american saurolophines unfortunately the nasal bones are also incomplete in the skull remains from this material 26 the informal name sabinosaurus has been given to the specimen 28 a possibly second but confirmed to be valid species of kritosaurus may have lived in the javelina formation alongside kritosaurus navajovius 30 31 32 33 kritosaurus fossils have also been unearthed in the olmos formation in mexico 34 description edit size comparison the type specimen of kritosaurus navajovius is only represented by a partial skull and lower jaws and associated postcranial remains 13 the greater portion of the muzzle and upper beak are missing 26 however these remains alone indicate a large body size reaching 9 metres 30 ft in length and 4 metric tons 4 4 short tons in body mass 35 36 the length of the skull is estimated at 87 cm 34 in from the tip of the upper beak to the base of the quadrate that articulates with the lower jaw at the back of the skull 37 based on the skull originally referred to anasazisaurus the form of the complete crest is that of a tab or flange of bone from the nasals that rises between and above the eyes and folds back under itself this unique crest allows it to be distinguished from similar hadrosaurs like gryposaurus 23 the top of the crest is roughened and the maximum preserved length of the skull could reach 90 cm 35 in 29 potential diagnostic characteristics of kritosaurus include a predentary lower beak without tooth like crenulations a sharp downward bend to the lower jaws near the beak and a heavy somewhat rectangular maxilla upper tooth bearing bone 26 k navajovius life restoration according to prieto márquez who re diagnosed this genus in 2013 kritosaurus can be distinguished based on the following characteristics the length of the dorsolateral margin of the maxilla is extensive the jugal features an orbital constriction that is deeper than the infratemporal one the infratemporal fenestra is greater than the orbit and has a dorsal margin that is greatly elevated above the dorsal orbital margin in adults the frontal bone is participating in the orbital margin the presence of paired caudal parasagittal processes of the nasals resting over the frontal bones 27 classification edit kritosaurus was a hadrosaurine hadrosaurid a flat headed or solid crested duckbill though many species and specimens have been referred to the genus in the past most of them do not show the shared distinguishing characteristics to allow them to be considered part of the genus or have been synonymized with other genera of hadrosaurs the closest relative of kritosaurus navajovius is anasazisaurus horneri or kritosaurus horneri which together with close relatives such as gryposaurus and secernosaurus form a clade called the kritosaurini within the larger clade saurolophinae 27 location and time separate kritosaurus and the slightly older primarily canadian gryposaurus along with some cranial details 26 cast of an assigned egg the following is a cladogram based on the phylogenetic analysis conducted by prieto márquez and wagner in 2012 showing the relationships of kritosaurus among the other kritosaurini 38 kritosaurini wulagasaurus kritosaurus gryposaurus latidens gryposaurus notabilis gryposaurus monumentensis aquilarhinus palimentus secernosaurus willinakaqe paleobiology edit the nasal crest of kritosaurus whatever its true form may have been used for a variety of social functions such as identification of sexes or species and social ranking 13 there may have been inflatable air sacs flanking it for both visual and auditory signaling 18 diet and feeding edit lower jaw in inner view as a hadrosaurid kritosaurus would have been a large bipedal quadrupedal herbivore eating plants with a sophisticated skull that permitted a grinding motion analogous to chewing its teeth were continually replacing and packed into dental batteries that contained hundreds of teeth only a relative handful of which were in use at any time plant material would have been cropped by its broad beak and held in the jaws by a cheek like organ feeding would have been from the ground up to 4 meters 13 ft above 13 if it was a separate genus how it would have partitioned resources with the similar and contemporaneous naashoibitosaurus is unknown citation needed paleoecology edit tooth museo civico di storia naturale di milano kritosaurus was discovered in the de na zin member of the kirtland formation this formation dates from the late campanian stages of the late cretaceous period 74 to 70 million years ago 4 and is also the source of several other dinosaurs like alamosaurus a species of parasaurolophus pentaceratops nodocephalosaurus saurornitholestes and bistahieversor 39 the kirtland formation is interpreted as river floodplains appearing after a retreat of the western interior seaway conifers were the dominant plants and chasmosaurine horned dinosaurs appear to have been more common than hadrosaurids 40 the presence of parasaurolophus and kritosaurus in northern latitude fossil sites may represent faunal exchange between otherwise distinct northern and southern biomes in late cretaceous north america 41 both taxa are uncommon outside of the southern biome where along with pentaceratops they are predominate members of the fauna 41 the geographic range of kritosaurus remains in north america was expanded by the discovery of bones from the late campanian age aguja formation of texas including a skull 42 32 although this specimen was given its own genus name aquilarhinus in 2019 43 additionally a partial skull from coahuila mexico has been referred to k navajovius 27 since the 1910s and 1930s barnum brown described that an unsubscribed species of kritosaurus the most likely candidate being kritosaurus navajovius had inhabited the late maastrichtian ojo alamo formation where the first specimen of kritosaurus was unearthed in new mexico as well as the javelina formation and the el picacho formation in texas which was a flood plain type environment at the time of the cretaceous 2 44 45 46 charles w gilmore also made notes about brown s work surveys and finds from the ojo alamo formation whilst doing research in the north horn formation in utah as well as researching the ojo alamo formation himself 47 48 these fossils might be of an unknown species of hadrosaur or an undescribed specimen of kritosaurus or kritosaurus navajovius however not all of the paleontological community agrees with the age of the kritosaurus holotype unearthed by barnum brown this is due to the unconformity that divides the ojo alamo formation into two parts the older naashoibito member which overlies the campanian era kirtland formation and the younger kimbeto member starting in the 2000s and 2010s more research into this area as well as nearby fossil formations in neighboring states has brought more information about them to light this issue will probably be resolved in the future 49 50 51 52 53 54 however confirmed kritosaurus remains possibly belonging to k navajovius cf k navajovius and possibly a new species have been unearthed in the javelina formation and the el picacho formation in texas 32 2 44 45 55 56 57 58 59 this genus lived alongside numerous species of dinosaurs including the sauropod alamosaurus the ceratopsians bravoceratops ojoceratops torosaurus and a possible species of eotriceratops hadrosaurs which included a possible species of edmontosaurus annectens a hadrosaur very similar to saurolophus and gryposaurus gryposaurus alsatei to be exact 55 and the armored nodosaur glyptodontopelta theropods from this environment which included tyrannosaurus smaller theropods like a species of troodon and richardoestesia the oviraptorid ojoraptorsaurus the dromaeosaur dineobellator and indeterminate ornithomimids and other undescribed dromaeosaurs non dinosaur species that had shared the same environment with kritosaurus included the giant pterosaur quetzalcoatlus various species of fishes and rays amphibians lizards turtles like adocus and multiple species of mammals like alphadon and mesodma see also edit timeline of hadrosaur research references edit creisler benjamin s 2007 deciphering duckbills in carpenter kenneth ed horns and beaks ceratopsian and ornithopod dinosaurs bloomington and indianapolis indiana university press pp 185 210 isbn 978 0 253 34817 3 1 2 3 4 brown barnum 1910 the cretaceous ojo alamo beds of new mexico with description of the new dinosaur genus kritosaurus bulletin of the american museum of natural history 28 24 267 274 hdl 2246 1398 1 2 gilmore charles w 1916 contributions to the geology and paleontology of san juan county new mexico 2 vertebrate faunas of the ojo alamo kirtland and fruitland formations united states geological survey professional paper 98 q 279 302 1 2 3 williamson thomas e 2000 review of hadrosauridae dinosauria ornithischia from the san juan basin new mexico in lucas s g heckert a b eds dinosaurs of new mexico new mexico museum of natural history and science bulletin 17 albuquerque new mexico new mexico museum of natural history and science pp 191 213 1 2 sinclair william j granger walter 1914 paleocene deposits of the san juan basin new mexico bulletin of the american museum of natural history 33 3 297 316 bibcode 1916jg 24q 305s doi 10 1086 622336 olshevsky george 1999 11 17 re what are these dinosaurs 2 return of what are these dinosaurs dinosaur mailing list archived from the original on 2011 11 17 retrieved 2007 06 15 lambe lawrence m 1914 on gryposaurus notabilis a new genus and species of trachodont dinosaur from the belly river formation of alberta with a description of the skull of chasmosaurus belli the ottawa naturalist 27 11 145 155 brown barnum 1914 cretaceous eocene correlation in new mexico wyoming montana alberta geological society of america bulletin 25 1 355 380 bibcode 1914gsab 25 355b doi 10 1130 gsab 25 355 hdl 2246 704 archived from the original on 2022 12 12 retrieved 2020 09 13 parks william a 1920 the osteology of the trachodont dinosaur kritosaurus incurvimanus university of toronto studies geology series 11 1 76 prieto marquez alberto 2010 the braincase and skull roof of gryposaurus notabilis dinosauria hadrosauridae with a taxonomic revision of the genus journal of vertebrate paleontology 30 3 838 854 bibcode 2010jvpal 30 838p doi 10 1080 02724631003762971 s2cid 83539808 1 2 lull richard swann wright nelda e 1942 hadrosaurian dinosaurs of north america geological society of america special paper 40 geological society of america pp 164 172 marsh o c 1889 notice of new american dinosauria american journal of science 38 220 331 336 bibcode 1889amjs 37 331m doi 10 2475 ajs s3 37 220 331 s2cid 131729220 archived from the original on 2022 12 16 retrieved 2019 09 11 1 2 3 4 5 horner john r weishampel david b forster catherine a 2004 hadrosauridae in weishampel david b dodson peter osmólska halszka eds the dinosauria 2nd ed berkeley university of california press pp 438 463 isbn 978 0 520 24209 8 prieto márquez alberto weishampel david b horner john r 2006 the dinosaur hadrosaurus foulkii from the campanian of the east coast of north america with a reevaluation of the genus pdf acta palaeontologica polonica 51 1 77 98 arc...
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