Hadrosauromorph
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Hadrosauromorpha is a clade of
iguanodontia Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
n
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
s, defined in
2014 The year 2014 was marked by the surge of the Western African Ebola epidemic, West African Ebola epidemic, which began in 2013, becoming the List of Ebola outbreaks, most widespread outbreak of the Ebola, Ebola virus in human history, resul ...
by David B. Norman to divide
Hadrosauroidea Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or Hadrosauridae, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Euro ...
into the basal taxa with compressed manual bones and a pollex, and the derived taxa that lack them. The clade is formally defined in 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 ...
'' as "the largest clade containing ''
Hadrosaurus foulkii ''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 in New Jersey about 83.6 to 77.9 Ma. The holotype specimen was found in fluv ...
'', but not ''
Probactrosaurus gobiensis ''Probactrosaurus'' (meaning "before ''Bactrosaurus''") is an early herbivorous hadrosauroid iguanodont dinosaur. It lived in China during the Early Cretaceous period. Discovery and species In 1959 and 1960 a Soviet-Chinese expedition uncovered ...
''". 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 (zoology), specific name honouring William Henry Fitton. History and n ...
''. 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 first given a phylogenetic definition by Norman (2014 and 2015) as
hadrosauroid Hadrosauroidea is a clade or superfamily of ornithischian dinosaurs that includes the "duck-billed" dinosaurs, or Hadrosauridae, and all dinosaurs more closely related to them than to ''Iguanodon''. Their remains have been recovered in Asia, Euro ...
taxa closer to ''
Edmontosaurus regalis ''Edmontosaurus regalis'' is a species of comb-crested hadrosaurid dinosaur. Fossils of ''E. regalis'' have been found in rocks of western North America that date from the late Campanian age of the Cretaceous Geologic time scale#Terminology, Peri ...
'' than ''
Probactrosaurus gobiensis ''Probactrosaurus'' (meaning "before ''Bactrosaurus''") is an early herbivorous hadrosauroid iguanodont dinosaur. It lived in China during the Early Cretaceous period. Discovery and species In 1959 and 1960 a Soviet-Chinese expedition uncovered ...
''. 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 in New Jersey about 83.6 to 77.9 Ma. The holotype specimen was found in flu ...
'' 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 (), also hadrosaurs 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 fam ...
, as well as the taxa ''
Tethyshadros ''Tethyshadros'' ("Tethys Ocean, Tethyan hadrosauroid") is a genus of Hadrosauroidea, hadrosauroid dinosaur from the Late Cretaceous (Campanian) Calcare di Aurisina (previously thought to come from the younger Liburnia Formation) of Trieste, Ita ...
'' 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, from about 96 to 85 million years ago. The position ''Bactrosaurus'' ...
''. Norman in 2014 had included more taxa in Hadrosauromorpha, those of Norman (2015) as well as '' Levnesovia'', ''
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 (phylogeny), 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 b ...
'', 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 19 ...
'' 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 (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
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, the crown is the visible part of the tooth above the gingival margin and is an essential component of dental anatomy. Covered by Tooth enamel, enamel, the crown plays a crucial role in cutting, tearing, and grinding food. Its shap ...
s in the
dentary In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone ...
are asymmetrical and have multiple vertical ridges; there is a
foramen In anatomy and osteology, a foramen (; : foramina, or foramens ; ) is an opening or enclosed gap within the dense connective tissue (bones and deep fasciae) of extant and extinct amniote animals, typically to allow passage of nerves, artery, ...
in the
surangular The surangular or suprangular 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 bone, angular, splenial and articular. It is o ...
; 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 up ...
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 mammals h ...
contacts the
prefrontal Prefrontal may refer to: *Prefrontal bone, a skull bone in some tetrapods *Prefrontal cortex, a region of the brain of a mammal *Prefrontal scales The prefrontal scales on snakes and other reptiles are the scales adjacent and anterior to the fr ...
, and the jugal contact with the ectopterygoid bone of the palate is reduced. In the appendicular regions, hadrosauromorphans the
scapula The scapula (: 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 either side ...
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 inst ...
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 found worldwide have been classified in the genus ''Iguanodon'', dating from the Late Jurassic to Early Cretaceous, Taxonomy (bi ...
'' 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 (zoology), specific name honouring William Henry Fitton. History and n ...
''. 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 (carpus) that connects the hand to the forearm. The terms "carpus" and "carpal" are derived from the Latin wikt:carpus#Latin, carpus and the Greek language, Greek wikt:καρπός ...
, and a less mobile
manus Manus may refer to: Relating to locations around New Guinea *Manus Island, a Papua New Guinean island in the Admiralty Archipelago ** Manus languages, languages spoken on Manus and islands close by ** Manus Regional Processing Centre, an offshore ...
. 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 Hadrosauroidea Late Cretaceous dinosaurs Dinosaur clades