Hadrosauromorpha
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Hadrosauromorpha is a clade of
iguanodontia Iguanodontia (the iguanodonts) is a clade of herbivorous dinosaurs that lived from the Middle Jurassic to Late Cretaceous. Some members include ''Camptosaurus'', ''Dryosaurus'', ''Iguanodon'', ''Tenontosaurus'', and the hadrosaurids or "duck-bi ...
n
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...
s, defined in
2014 File:2014 Events Collage.png, From top left, clockwise: Stocking up supplies and personal protective equipment (PPE) for the Western African Ebola virus epidemic; Citizens examining the ruins after the Chibok schoolgirls kidnapping; Bundles of wat ...
by David B. Norman to divide
Hadrosauroidea Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or hadrosaurids, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Europ ...
into the basal taxa with compressed manual bones and a pollex, and the derived taxa that lack them. The clade is defined as all the taxa closer to ''
Edmontosaurus regalis ''Edmontosaurus regalis'' is a species of comb-crested hadrosaurid (duck-billed) dinosaur. Fossils of ''E. regalis'' have been found in rocks of western North America that date from the late Campanian stage of the Cretaceous Period 73 million yea ...
'' than '' Probactrosaurus gobiensis''. This results in different taxon inclusion depending on the analysis.


Classification

Hadrosauromorpha was first used in literature by David B. Norman in 2014 in a discussion of phylogenetics of ''
Hypselospinus ''Hypselospinus'' is a genus of iguanodontian dinosaur which was first described as a species of ''Iguanodon'' (''I. fittoni'') by Richard Lydekker in 1889, the specific name honouring William Henry Fitton. History In May 2010 the fossils com ...
''. In his 2014 paper Norman references another of his publications as the authority for Hadrosauromorpha, a chapter in the book ''Hadrosaurs''. However, the book was in fact published later, in 2015. Following Article 19.4 of the
PhyloCode The ''International Code of Phylogenetic Nomenclature'', known as the ''PhyloCode'' for short, is a formal set of rules governing phylogenetic nomenclature. Its current version is specifically designed to regulate the naming of clades, leaving the ...
, the authorship of the clade is thus Norman (2015), while the authorship of the definition is Norman (2014).


Definition

Hadrosauromorpha was defined by Norman (2014 and 2015) as
hadrosauroid Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or hadrosaurids, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Eu ...
taxa closer to ''
Edmontosaurus regalis ''Edmontosaurus regalis'' is a species of comb-crested hadrosaurid (duck-billed) dinosaur. Fossils of ''E. regalis'' have been found in rocks of western North America that date from the late Campanian stage of the Cretaceous Period 73 million yea ...
'' than '' Probactrosaurus gobiensis''. This definition was contested by Mickey Mortimer, who stated that to follow the PhyloCode the taxon ''
Hadrosaurus ''Hadrosaurus'' (; ) is a genus of hadrosaurid ornithopod dinosaurs that lived in North America during the Late Cretaceous Period in what is now the Woodbury Formation about 80 million to 78 million years ago. The holotype specimen was found in f ...
'' must be included in the definition, as it is the type genus of Hadrosauromorpha. By this definition, Norman (2015) considered Hadrosauromorpha to include
Hadrosauridae Hadrosaurids (), or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod family, which includ ...
, as well as the taxa ''
Tethyshadros ''Tethyshadros'' ("Tethys Ocean, Tethyan hadrosauroid") is a genus of hadrosauroidea, hadrosauroid dinosaur from Trieste, Italy. The type and only species is ''T. insularis''. Discovery and naming Sometime in the 1980s, Alceo Tarlao and Giorgi ...
'' and ''
Bactrosaurus ''Bactrosaurus'' (; meaning "Club lizard," "baktron" = club + ''sauros'' = lizard) is a genus of herbivorous hadrosauroid dinosaur that lived in Asia during the Late Cretaceous, about 96-85 million years ago. The position ''Bactrosaurus'' occupie ...
''. Norman in 2014 had included more taxa in Hadrosauromorpha, those of Norman (2015) as well as ''
Levnesovia ''Levnesovia'' is a genus of hadrosauroid dinosaur from the Late Cretaceous Bissekty Formation of Uzbekistan. It was related to ''Bactrosaurus''. The type species is ''L. transoxiana''. The genus name honours the late Russian paleontologist Lev ...
'', ''
Gilmoreosaurus ''Gilmoreosaurus'' (meaning "Charles Whitney Gilmore's lizard") is the name given to a genus of dinosaur from the Cretaceous of Asia. The type species is ''Gilmoreosaurus mongoliensis''. It is believed to be a hadrosaur or iguanodont from the Ire ...
'' and ''
Telmatosaurus ''Telmatosaurus'' (meaning "marsh lizard") is a genus of basal hadrosauromorph dinosaur from the Late Cretaceous of Romania. It was a relatively small hadrosaur, measuring approximately in length and in body mass, which has been explained as a ...
'', the last of which was considered inside Hadrosauridae by Norman in 2015. Another phylogenetic analysis by Xing ''et al.'' in 2014 also found that ''
Eolambia ''Eolambia'' (meaning "dawn lambeosaurine") is a genus of herbivorous hadrosauroid dinosaur from the early Late Cretaceous of the United States. It contains a single species, ''E. caroljonesa'', named by paleontologist James Kirkland in 1998. ...
'' and ''
Protohadros ''Protohadros'' (meaning "first hadrosaur") is a genus of herbivorous ornithischian dinosaur from the Late Cretaceous ( Cenomanian stage). Gary Byrd, a part-time palaeontologist, discovered some remains of this euornithopod (ribs and an ungual) ...
'', both found outside Hadrosauromorpha by Norman, fell within his definition, as well as a large number of other taxa.


Phylogeny

Many different versions of phylogenies have been conducted on the group of hadrosauromorphs. Norman (2014) created his own analysis, which includes 105 different morphological characters and 27 select
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...
taxa. His phylogeny is shown below, using his specific clade definitions: Norman's definitions have been heavily criticized by Mickey Mortimer as being unnecessary changes which cause more confusion to classification. Other phylogenetic analyses, like the results of Madzia ''et al.'' in 2020, have placed Hadrosauromorpha in the middle of a long grade of stem-hadrosaurs, without any large groups of taxa unlike previous versions of the same analysis, apart from a large group of Eurasian taxa.


Description

''Probactrosaurus'' was selected as the outgroup to Hadrosauromorpha because of numerous differences that Norman (2014) thought to be significant. The
tooth crown In dentistry, crown refers to the anatomical area of teeth, usually covered by tooth enamel, enamel. The crown is usually visible in the mouth after tooth development, developing below the gingiva and then tooth eruption, erupting into place. ...
s in the
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower tooth, teeth in place. The mandible sits beneath the maxilla. It is the only movabl ...
are asymmetrical and have multiple vertical ridges; there is a
foramen In anatomy and osteology, a foramen (;Entry "foramen"
in
in the
surangular The suprangular or surangular is a jaw bone found in most land vertebrates, except mammals. Usually in the back of the jaw, on the upper edge, it is connected to all other jaw bones: dentary, angular, splenial and articular The articular bone i ...
; and the
quadrate bone The quadrate bone is a skull bone in most tetrapods, including amphibians, sauropsids (reptiles, birds), and early synapsids. In most tetrapods, the quadrate bone connects to the quadratojugal and squamosal bones in the skull, and forms upper ...
has a more prominent depression for the articulation of the
jugal The jugal is a skull bone found in most reptiles, amphibians and birds. In mammals, the jugal is often called the malar or zygomatic. It is connected to the quadratojugal and maxilla, as well as other bones, which may vary by species. Anatomy ...
. None of these features are found in the skulls of the more derived hadrosauromorphans. The
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has b ...
contacts the prefrontal, and the jugal contact with the ectopterygoid bone of the palate is reduced. In the appendicular regions, hadrosauromorphans the
scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...
e are not J shaped, instead having an overhanging projection. The lower
forelimb A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial ( anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...
bones are more slender in both hadrosauromorphans and ''Probactosaurus'', unlike their more robust ancestors. ''Probactosaurus'', however, possesses the basal condition of having a small, conical pollex, like in earlier ornithopods such as ''
Iguanodon ''Iguanodon'' ( ; meaning 'iguana-tooth'), named in 1825, is a genus of iguanodontian dinosaur. While many species have been classified in the genus ''Iguanodon'', dating from the late Jurassic Period to the early Cretaceous Period of Asia, Eu ...
'' or ''
Hypselospinus ''Hypselospinus'' is a genus of iguanodontian dinosaur which was first described as a species of ''Iguanodon'' (''I. fittoni'') by Richard Lydekker in 1889, the specific name honouring William Henry Fitton. History In May 2010 the fossils com ...
''. This absence of a pollex is also linked to a reduction of the
carpal bones The carpal bones are the eight small bones that make up the wrist (or carpus) that connects the hand to the forearm. The term "carpus" is derived from the Latin carpus and the Greek καρπός (karpós), meaning "wrist". In human anatomy, th ...
, and a less mobile manus. The ilium bones of ''Probactrosaurus'' are more angular than in hadrosauromorphs, which lack a brevis shelf. It was also identified that the femoral shaft is straight in hadrosauromorphans and the pedal bones are truncated


References

{{Taxonbar, from=Q48997072 Iguanodonts Cretaceous dinosaurs Taxa named by David B. Norman