''Acristavus'' (meaning "non-crested grandfather") is a

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

saurolophine Saurolophinae is a subfamily (biology), subfamily of hadrosaurid dinosaurs. It has since the mid-20th century generally been called the Hadrosaurinae, a group of largely non-crested hadrosaurs related to the crested sub-family Lambeosaurinae. How ...

dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the evolution of dinosaurs is t ...

. Fossils have been found from the

Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campanian s ...

Two Medicine Formation The Two Medicine Formation is a geological formation, or rock body, in northwestern Montana and southern Alberta that was deposited between and (million years ago), during Campanian (Late Cretaceous) time. It crops out to the east of the Rocky M ...

Montana Montana () is a state in the Mountain West division of the Western United States. It is bordered by Idaho to the west, North Dakota and South Dakota to the east, Wyoming to the south, and the Canadian provinces of Alberta, British Columbi ...

and

Wahweap Formation The Wahweap Formation of the Grand Staircase–Escalante National Monument is a geological formation in southern Utah and northern Arizona, around the Lake Powell region, whose strata date back to the Late Cretaceous (Campanian stage). Dinosaur r ...

Utah Utah ( , ) is a state in the Mountain West subregion of the Western United States. Utah is a landlocked U.S. state bordered to its east by Colorado, to its northeast by Wyoming, to its north by Idaho, to its south by Arizona, and to it ...

United States The United States of America (U.S.A. or USA), commonly known as the United States (U.S. or US) or America, is a country primarily located in North America. It consists of 50 states, a federal district, five major unincorporated territorie ...

. The

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

''A. gagslarsoni'' was named in 2011. Unlike nearly all hadrosaurids except ''

Edmontosaurus ''Edmontosaurus'' ( ) (meaning "lizard from Edmonton") is a genus of hadrosaurid (duck-billed) dinosaur. It contains two known species: ''Edmontosaurus regalis'' and ''Edmontosaurus annectens''. Fossils of ''E. regalis'' have been found in rocks ...

'', ''Acristavus'' lacked ornamentation on its skull. The discovery of ''Acristavus'' is paleontologically significant because it supports the position that the ancestor of all hadrosaurids did not possess cranial ornamentation, and that ornamentation was an adaptation that later arose interdependently in the subfamilies Saurolophinae and

Lambeosaurinae Lambeosaurinae is a group of crested hadrosaurid dinosaurs. Classification Lambeosaurines have been traditionally split into the tribes or clades Parasaurolophini (''Parasaurolophus'', ''Charonosaurus'', others (?).) and Lambeosaurini (''Coryt ...

. It is closely related to ''

Brachylophosaurus ''Brachylophosaurus'' ( or ; meaning "short-crested lizard", Greek ''brachys'' = short + ''lophos'' = crest + ''sauros'' = lizard, referring to its small crest) was a mid-sized member of the hadrosaurid family of dinosaurs. It is known from sev ...

'' and ''

Maiasaura ''Maiasaura'' (from the Greek ''μαῖα'', meaning "good mother" and ''σαύρα'', the feminine form of ''saurus'', meaning "reptile") is a large herbivorous saurolophine hadrosaurid ("duck-billed") dinosaur genus that lived in the area curre ...

'', and was assigned to a new

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

called

Brachylophosaurini Brachylophosaurini is a tribe of saurolophine hadrosaurs with known material being from N. America and potentially Asia. It contains at least four taxa; ''Acristavus'' (from Montana and Utah), ''Brachylophosaurus'' (from Montana and Alberta), '' ...

Discovery and occurrence

left, Skull The

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

specimen of ''Acristavus'', MOR 1155, was recovered at the Two Medicine Formation, in Teton County, Montana. The specimen was collected in 1999 by C. Riley Nelson in well-

indurated Friability ( ), the condition of being friable, describes the tendency of a solid substance to break into smaller pieces under duress or contact, especially by rubbing. The opposite of friable is indurate. Substances that are designated hazardous, ...

tan colored calcareous sandstone that was deposited during the

stage Stage or stages may refer to: Acting * Stage (theatre), a space for the performance of theatrical productions * Theatre, a branch of the performing arts, often referred to as "the stage" * ''The Stage'', a weekly British theatre newspaper * Sta ...

of the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...

period, approximately 79 million years ago. MOR 1155 consists of an almost complete skull with associated postcrania including eleven cervical vertebrae, three incomplete dorsal vertebrae, a proximal caudal vertebra, several dorsal ribs, the left humerus, the left ulna, the right sternal, the left pubis, the left femur, the left tibia, two left metatarsals, five left pedal phalanges and one right pedal phalanx. A second specimen UMNHVP 16607 assigned to ''Acristavus'' in 2011, was excavated from the Smokey Mountain Road locality in the Reynolds Point Member of the Wahweap Formation in Utah. It was collected by C. R. Nelson in 2000, in lithified, yellow sandstone and its age was estimated to be 80.69 Ma, with a range of uncertainty of 81.05-80.19 Ma. UMNHVP 16607 consists of a partial articulated skull, including both lacrimals, a complete braincase, and a cervical vertebra.

Classification

''Acristavus'' was first assigned to the

, in a basal position, by Gates et al. (2011). All subsequent

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

analyses have confirmed this assignment. Brachylophosaurines are derived members of the group

Saurolophinae Saurolophinae is a subfamily of hadrosaurid dinosaurs. It has since the mid-20th century generally been called the Hadrosaurinae, a group of largely non-crested hadrosaurs related to the crested sub-family Lambeosaurinae. However, the name Hadro ...

. Other brachylophosaurins include ''

'', ''

'', and potentially ''

Wulagasaurus ''Wulagasaurus'' (meaning "Wulaga lizard", in reference to the discovery locality) is a genus of saurolophine hadrosaurid dinosaur from the Late Cretaceous of Heilongjiang, China. Its remains were found in a bonebed in the middle Maastrichtian- ...

''. Gates concluded that ''Acristavus'' and ''Maiasaura'' shared a sister-

taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

relationship but more recent analysis by Prieto-Márquez (2013) shows that ''Maiasaura'' is more closely related to the more derived ''Brachylophosaurus''. The following cladogram is based on the 2013 phylogenetic analysis by Prieto-Márquez (the relationships within Lambeosaurinae and between basal hadrosauroids are not shown):

Distinguishing anatomical features

A diagnosis is a statement of the anatomical features of an organism (or group) that collectively distinguish it from all other organisms. Some, but not all, of the features in a diagnosis are also autapomorphies. An autapomorphy is a distinctive anatomical feature that is unique to a given organism. According to Gates (2011), ''Acristavus'' can be distinguished based on the following characteristics: * an enlarged caudodorsal brow of the postorbital and the basioccipital, which extends caudally well beyond the dorsal border of the foremen magnum to a greater extent than in other hadrosaurine species * a deep laterally squared frontonasal suture * swelling of the interdigitate suture between the prefrontal and frontal * a deep depression is present on the lateral surface of the postorbital jugal process

Paleoecology

Habitat

The Wahweap Formation has been radiometrically dated as being between 81 and 76 million years old. During the time that ''Acristavus'' lived, the

Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea, ...

was at its widest extent, almost completely isolating southern

Laramidia Laramidia was an island continent that existed during the Late Cretaceous period (99.6–66 Ma), when the Western Interior Seaway split the continent of North America in two. In the Mesozoic era, Laramidia was an island land mass separated from A ...

off from the rest of North America. The area where dinosaurs lived included lakes, floodplains, and east-flowing rivers. The Wahweap Formation is part of the

Grand Staircase The Grand Staircase is an immense sequence of sedimentary rock layers that stretch south from Bryce Canyon National Park and Grand Staircase–Escalante National Monument, through Zion National Park, and into Grand Canyon National Park. Ch ...

region, an immense sequence of sedimentary rock layers that stretch south from Bryce Canyon National Park through Zion National Park and into the Grand Canyon. The presence of rapid sedimentation and other evidence suggests a wet, seasonal climate.

Paleofauna

''Acristavus'' shared its

paleoenvironment Paleoecology (also spelled palaeoecology) is the study of interactions between organisms and/or interactions between organisms and their environments across geologic timescales. As a discipline, paleoecology interacts with, depends on and informs ...

with other dinosaurs, such as the

lambeosaur Lambeosaurinae is a group of crested hadrosaurid dinosaurs. Classification Lambeosaurines have been traditionally split into the tribes or clades Parasaurolophini (''Parasaurolophus'', ''Charonosaurus'', others (?).) and Lambeosaurini (''Coryt ...

'' Adelolophus hutchisoni'', the

ceratopsian Ceratopsia or Ceratopia ( or ; Greek: "horned faces") is a group of herbivorous, beaked dinosaurs that thrived in what are now North America, Europe, and Asia, during the Cretaceous Period, although ancestral forms lived earlier, in the Jurassic. ...

Diabloceratops ''Diabloceratops'' is an extinct genus of centrosaurine ceratopsian dinosaur that lived approximately 81.4-81 million years ago during the latter part of the Cretaceous Period in what is now Utah, in the United States. ''Diabloceratops'' was a ...

eatoni'', unnamed ankylosaurs and pachycephalosaurs, and the

theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...

Lythronax ''Lythronax'' () is a genus of tyrannosaurid dinosaur that lived in North America around 81.9-81.5 million years ago during the Late Cretaceous period. The only known specimen was discovered in Utah in the Wahweap Formation of the Grand Stairca ...

argestes'', which was likely the

apex predator An apex predator, also known as a top predator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the highest trophic lev ...

in its ecosystem. Vertebrates present in the Wahweap Formation at the time of ''Acristavus'' included

freshwater fish Freshwater fish are those that spend some or all of their lives in fresh water, such as rivers and lakes, with a salinity of less than 1.05%. These environments differ from marine conditions in many ways, especially the difference in levels of s ...

bowfin The bowfin (''Amia calva'') is a bony fish, native to North America. Common names include mudfish, mud pike, dogfish, grindle, grinnel, swamp trout, and choupique. It is regarded as a relict, being the sole surviving species of the Halecomorphi ...

s, abundant

ray Ray may refer to: Fish * Ray (fish), any cartilaginous fish of the superorder Batoidea * Ray (fish fin anatomy), a bony or horny spine on a fin Science and mathematics * Ray (geometry), half of a line proceeding from an initial point * Ray (g ...

s and

sharks Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorp ...

, turtles like ''

Compsemys ''Compsemys '' is an extinct genus of prehistoric turtles from the Late Cretaceous and Paleocene of North America and possibly Europe. The type species ''C. victa'', first described by Joseph Leidy from the Hell Creek Formation in Montana in 1856 ...

'', crocodilians, and

lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...

. A fair number of mammals lived in this region, which included several genera of

multituberculates Multituberculata (commonly known as multituberculates, named for the multiple tubercles of their teeth) is an extinct order of rodent-like mammals with a fossil record spanning over 130 million years. They first appeared in the Middle Jurassic, ...

cladotheria Cladotheria is a clade (sometimes ranked as a legion) of mammals. It contains modern therian mammals (marsupials and placentals) and several extinct groups, such as the dryolestoids, amphitheriids and peramurids. The clade was named in 1975 by M ...

ns,

marsupials Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsupials are endemic to Australasia, Wallacea and the Americas. A distinctive characteristic common to most of these species is that the young are carried in a po ...

, and

placental Placental mammals (infraclass Placentalia ) are one of the three extant subdivisions of the class Mammalia, the other two being Monotremata and Marsupial Marsupials are any members of the mammalian infraclass Marsupialia. All extant marsup ...

insectivores A robber fly eating a hoverfly An insectivore is a carnivorous animal or plant that eats insects. An alternative term is entomophage, which can also refer to the human practice of eating insects. The first vertebrate insectivores wer ...

. These mammals were more primitive than those that lived in the area that is now the

Kaiparowits Formation The Kaiparowits Formation is a sedimentary rock formation found in the Kaiparowits Plateau in Grand Staircase–Escalante National Monument, in the southern part of Utah in the western United States. It is over 2800 feet (850 m) thick, and is Ca ...

Trace fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or animal itself. Trace fossils contrast with body fossils, ...

s are relatively abundant in the Wahweap Formation, and suggest the presence of

crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...

, as well as

ornithischian Ornithischia () is an extinct order of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek s ...

and

dinosaurs. In 2010 a unique trace fossil was discovered that suggests a predator-prey relationship between dinosaurs and primitive mammals. The trace fossil includes at least two fossilized mammalian den complexes as well as associated digging grooves presumably caused by a

maniraptora Maniraptora is a clade of coelurosaurian dinosaurs which includes the birds and the non-avian dinosaurs that were more closely related to them than to ''Ornithomimus velox''. It contains the major subgroups Avialae, Deinonychosauria, Oviraptoros ...

n dinosaur. The proximity indicates a case of probable active predation of the burrow inhabitants by the animals that made the claw marks. Invertebrate activity in this formation ranged from fossilized

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...

burrows in petrified logs to various

mollusk Mollusca is the second-largest phylum of invertebrate animals after the Arthropoda, the members of which are known as molluscs or mollusks (). Around 85,000 extant species of molluscs are recognized. The number of fossil species is e ...

s, large

crab Crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" (abdomen) ( el, βραχύς , translit=brachys = short, / = tail), usually hidden entirely under the thorax. They live in all the ...

s,Kirkland, James Ian. "An inventory of paleontological resources in the lower Wahweap Formation (lower Campanian), southern Kaiparowits Plateau, Grand Staircase-Escalante National Monument, Utah." Abstracts with Programs - Geological Society of America, vol.37, no.7, pp.114, Oct 2005. and a wide diversity of

gastropod The gastropods (), commonly known as snails and slugs, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, from freshwater, and from land. T ...

s and

ostracods Ostracods, or ostracodes, are a class of the Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 70,000 species (only 13,000 of which are extant) have been identified, grouped into several orders. They are small crustaceans, typical ...

.Williams, Jessica A J; Lohrengel, C Frederick. ''Preliminary study of freshwater gastropods in the Wahweap Formation, Bryce Canyon National Park, Utah.'' Abstracts with Programs - Geological Society of America, vol. 39, no. 5, pp.43, May 2007

References

{{Taxonbar, from=Q1932513 Saurolophines Late Cretaceous dinosaurs of North America Fossil taxa described in 2011 Taxa named by Jack Horner Paleontology in Montana Paleontology in Utah Campanian genus first appearances Campanian genus extinctions Ornithischian genera