Hadrosaurus
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''Hadrosaurus'' (; ) is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
hadrosaurid 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 inclu ...
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 w ...
dinosaurs 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 the ...
that lived in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...
during the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
Period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...
in what is now the
Woodbury Formation The Woodbury Formation is a Mesozoic geologic formation in New Jersey, United States. It primarily consists of massive dark gray clays and silts with carbonized wood fragments and pyrite, deposited in a marine setting. The remains of the dinosaur ...
about 80 million to 78 million years ago. 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 seve ...
specimen was found in fluvial marine sedimentation, meaning that the corpse of the animal was transported by a river and washed out to sea. They were large animals ranging from and . Most of the preserved elements are very robust, unusual traits in hadrosaurs. ''Hadrosaurus'' were ponderously-built animals equipped with keratinous beaks for cropping foliage and a specialized and complex dentition for food processing. ''Hadrosaurus foulkii'', the only
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...
in this
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
, is known from a single specimen consisting of much of the skeleton and parts of the skull. The specimen was collected in 1858 from the
Woodbury Formation The Woodbury Formation is a Mesozoic geologic formation in New Jersey, United States. It primarily consists of massive dark gray clays and silts with carbonized wood fragments and pyrite, deposited in a marine setting. The remains of the dinosaur ...
in
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delawa ...
, US, representing the first dinosaur species known from more than isolated teeth to be identified in North America. Using
radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares ...
of
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of marine and freshwater Mollusca, molluscs that have laterally compressed bodies enclosed by a shell consisting of two hing ...
shells from the same formation, the sedimentary rocks where the ''Hadrosaurus'' fossil was found have been
dated Date or dates may refer to: * Date (fruit), the fruit of the date palm (''Phoenix dactylifera'') Social activity * Dating, a form of courtship involving social activity, with the aim of assessing a potential partner **Group dating *Play date, ...
at some time between 80.5 and 78.5 million years ago. In 1858, the only known specimen became the first-ever dinosaur skeleton to be mounted. In 1991, ''H. foulkii'' became the official
state dinosaur State may refer to: Arts, entertainment, and media Literature * ''State Magazine'', a monthly magazine published by the U.S. Department of State * ''The State'' (newspaper), a daily newspaper in Columbia, South Carolina, United States * '' Our ...
of
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delawa ...
.


History of discovery

In 1838, John Estaugh Hopkins was digging in a
marl Marl is an earthy material rich in carbonate minerals, clays, and silt. When hardened into rock, this becomes marlstone. It is formed in marine or freshwater environments, often through the activities of algae. Marl makes up the lower part ...
pit (on a small tributary of the Cooper River in
Haddonfield, New Jersey :''Not the fictional Illinois town from the Halloween film series.'' Haddonfield is a borough located in Camden County, New Jersey, United States. As of the 2010 United States Census, the borough had a total population of 11,593,
, and part of 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. Campani ...
-age
Woodbury Formation The Woodbury Formation is a Mesozoic geologic formation in New Jersey, United States. It primarily consists of massive dark gray clays and silts with carbonized wood fragments and pyrite, deposited in a marine setting. The remains of the dinosaur ...
) when he uncovered large bones. He put them on display at his home, also in Haddonfield. In 1858, the bones sparked the interest of a visitor,
William Parker Foulke William Parker Foulke (1816–1865) discovered the first full dinosaur skeleton in North America (''Hadrosaurus foulkii'', which means "Foulke's big lizard") in Haddonfield, New Jersey in 1858. Born in Philadelphia, and a descendant of Welsh ...
, who dug out the full skeleton from the marl pit in the same year. The excavation site, known as the ''Hadrosaurus foulkii'' Leidy site, is now a
National Historic Landmark A National Historic Landmark (NHL) is a building, district, object, site, or structure that is officially recognized by the United States government for its outstanding historical significance. Only some 2,500 (~3%) of over 90,000 places liste ...
. Foulke contacted
paleontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of fossi ...
Joseph Leidy Joseph Mellick Leidy (September 9, 1823 – April 30, 1891) was an American paleontologist, parasitologist and anatomist. Leidy was professor of anatomy at the University of Pennsylvania, later was a professor of natural history at Swarthmore ...
, and together they recovered eight
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, ...
from the
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. T ...
r and
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 teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
areas, dental battery fragments, left maxilla fragments, three partial
dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...
, 13 caudal
centra Centra is a convenience shop chain that operates throughout Ireland. The chain operates as a symbol group owned by Musgrave Group, the food wholesaler, meaning the stores are all owned by individual franchisees. The chain has three different ...
, including an almost complete middle
caudal vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates,Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristic ...
. Other fragments included a partial right
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
, left
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a r ...
, left
radius In classical geometry, a radius (plural, : radii) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', ...
, left
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
, left ilium, right
ischium The ischium () form ...
, right partial pubis, the left hindlimb composed by the
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...
,
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
,
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
with
metatarsals The metatarsal bones, or metatarsus, are a group of five long bones in the foot, located between the tarsal bones of the hind- and mid-foot and the phalanges of the toes. Lacking individual names, the metatarsal bones are numbered from the me ...
II and IV and the first pedal
phalanx The phalanx ( grc, φάλαγξ; plural phalanxes or phalanges, , ) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar pole weapons. The term is particularly ...
from the third digit. Foulke and Leidy studied the fossils together and, in 1858, Leidy formally described and named ''Hadrosaurus foulkii'' in honor of his collaborator. While originally a portmanteau of Haddonfield, the location of its discovery with the accepted suffix for dinosaurs -saurus, the name ''Hadrosaurus'' was scientifically justified as deriving from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
, , meaning "bulky" or "large", and , , meaning "lizard". The name was an additional play on words by Leidy since it translates from Greek as ''Foulke’s big lizard''. Leidy recognized that the bones were from a dinosaur because of their similarity to those of ''
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, ...
'', discovered in England some decades before but, at the time, the skeleton of ''Hadrosaurus'' was one of the most complete dinosaur skeletons known. Leidy's monograph ''Cretaceous Reptiles of the United States'', describing ''Hadrosaurus'' more completely, and with illustrations, was written in 1860, but the
American Civil War The American Civil War (April 12, 1861 – May 26, 1865; also known by Names of the American Civil War, other names) was a civil war in the United States. It was fought between the Union (American Civil War), Union ("the North") and t ...
delayed its publication until 1865. Leidy reconstructed ''Hadrosaurus'' as a biped, in contrast to the view at the time that such dinosaurs were
quadruped Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
al. The entire skeleton was completely assembled in 1868 by a team including English
sculptor Sculpture is the branch of the visual arts that operates in three dimensions. Sculpture is the three-dimensional art work which is physically presented in the dimensions of height, width and depth. It is one of the plastic arts. Durable sc ...
and naturalist Benjamin Waterhouse Hawkins and was put on display at
Philadelphia Academy of Natural Sciences The Academy of Natural Sciences of Drexel University, formerly the Academy of Natural Sciences of Philadelphia, is the oldest natural science research institution and museum in the Americas. It was founded in 1812, by many of the leading natura ...
. It was the first-ever mounted dinosaur skeleton. When the skeleton was first put together, it was displayed with a plaster skull sculpted by Hawkins. Many other artists have recreated ''Hadrosaurus'' with skulls from other, related species such as '' Gryposaurus'' and ''
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 ...
''. A statue of ''Hadrosaurus'', sculpted by Haddonfield resident John Giannotti, now stands in the center of the town of Haddonfield, commemorating its discovery there. Thanks to Joyce Berry and her 4th grade classes (1988-1991) at Strawbridge Elementary School in Haddon Twp, NJ the ''Hadrosaurus'' was named the state dinosaur of
New Jersey New Jersey is a state in the Mid-Atlantic and Northeastern regions of the United States. It is bordered on the north and east by the state of New York; on the east, southeast, and south by the Atlantic Ocean; on the west by the Delawa ...
in 1991. It is one of the most celebrated dinosaurs ever and is of great historic importance. The skeleton is usually kept out of sight in the Academy's collections. However, from November 22, 2008, to April 19, 2009, a fully assembled cast of the skeleton, and an exhibit about the science and culture surrounding the dinosaur's discovery, was open to the public.


Description

''Hadrosaurus'' were large sized animals growing up to and weighing as much as .Genus List for Holtz 2012
/ref> According to Prieto-Márquez, ''Hadrosaurus'' can be distinguished in having a shortened pectoral crest that is slightly over 40% of the total humeral length, a deltopectoral crest that is developed from the humeral shaft causing the laterodistal border to display a broad lateral facet, a lower greatest point of the supraacetabular crest located above lateral edge from the rear to the bottom on the posterior tuberosity of the ischial peduncle of the ilium, a shortened supraacetabular crest from the front to the rear with its breadth being half the length of the middle iliac plate. As in most hadrosaurs, the forelimbs were not as heavily built as the hindlimbs, but were long enough to be used in standing or movement. The holotype specimen was a relatively large animal at the time of death with a long
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates ...
and long
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it conn ...
. Most of the preserved elements feature a marked robust composition with the
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
being one of the most robust among hadrosaurs.


Classification

Despite the fact that the family
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 inclu ...
has ''Hadrosaurus'' as its
type genus In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name. Zoological nomenclature According to the International Code of Zoological Nomenclature, "The name-bearing type of a nominal ...
, the skeleton lacks a skull and was long viewed as too incomplete to compare to other hadrosaurs for classification purposes, leading most scientists to consider it a ''nomen dubium'', or dubious name. However, a re-evaluation of the fossil material in 2011 noted several distinct characteristics of the skeleton that could allow the genus ''Hadrosaurus'' and species ''H. foulkii'' to remain in use as valid
taxa 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 ...
. ''Hadrosaurus'' has also traditionally served as the basis for a large subfamily called Hadrosaurinae, which was seen as a group of largely crestless hadrosaurs related to the crested subfamily Lambeosaurinae. However, the changing view of ''Hadrosaurus'' classification in relation to other hadrosaurs has led some scientists to rename these subfamilies. In a 2008 study, ''Hadrosaurus'' was found to be more primitive than either lambeosaurines or other "hadrosaurines", and not a particularly close relative of classic "hadrosaurines" such as ''
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 rock ...
'' and ''
Saurolophus ''Saurolophus'' (; meaning "lizard crest") is a genus of large hadrosaurid dinosaur from the Late Cretaceous period of Asia and North America, that lived in what is now the Horseshoe Canyon and Nemegt formations about 70 million to 68 million ...
''. As a result of this, the name Hadrosaurinae was restricted to ''Hadrosaurus'' alone, and the subfamily comprising the traditional "hadrosaurines" was renamed the
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 Hadr ...
. Below is a simplified
cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...
recovered by Ramírez-Velasco ''et al.'' in 2012 in their description of '' Huehuecanauhtlus''. This topology was recovered using an extensive sampling of 60 hadrosauroid species, and two outgroup taxa, which were scored based on 287 morphological traits, and included data from two recent redescriptions of ''Hadrosaurus'' by Prieto-Márquez ''et al.'' (2006) and Prieto-Márquez (2011).


Paleobiology

In 2003, Rothschild and colleagues performed a study looking for epidemiology of tumors in dinosaurs. Evidence of tumors, including hemangiomas, desmoplastic fibroma,
metastatic cancer Metastasis is a pathogenic agent's spread from an initial or primary site to a different or secondary site within the host's body; the term is typically used when referring to metastasis by a cancerous tumor. The newly pathological sites, the ...
, and osteoblastoma were discovered in specimens of ''Hadrosaurus'' by analyzing dinosaur vertebrae using computerized tomography and
fluoroscope Fluoroscopy () is an imaging technique that uses X-rays to obtain real-time moving images of the interior of an object. In its primary application of medical imaging Medical imaging is the technique and process of imaging the interior of a ...
screening. Several other hadrosaurids, including ''
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 ...
'', ''
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 rock ...
'', and '' Gilmoreosaurus'', also tested positive. Although more than 10000
fossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
were examined in this manner, the tumors were limited to ''Hadrosaurus'' and other hadrosaurs. The tumors were only found on caudal vertebrae and they may have been caused by environmental factors or genetic inheritance.


Paleoecology

The holotype of ''Hadrosaurus'' was found in marine sediments, which suggests the skeleton was transported by a river and then deposited in the Cretaceous sea. The ''Hadrosaurus'' remains all persist to the
Woodbury Formation The Woodbury Formation is a Mesozoic geologic formation in New Jersey, United States. It primarily consists of massive dark gray clays and silts with carbonized wood fragments and pyrite, deposited in a marine setting. The remains of the dinosaur ...
. Through the
radiometric dating Radiometric dating, radioactive dating or radioisotope dating is a technique which is used to date materials such as rocks or carbon, in which trace radioactive impurities were selectively incorporated when they were formed. The method compares ...
of
bivalve Bivalvia (), in previous centuries referred to as the Lamellibranchiata and Pelecypoda, is a class (biology), class of marine and freshwater Mollusca, molluscs that have laterally compressed bodies enclosed by a shell consisting of two hing ...
shells known from the formation, the sedimentary rocks where the holotype specimen of ''Hadrosaurus'' was found have been
dated Date or dates may refer to: * Date (fruit), the fruit of the date palm (''Phoenix dactylifera'') Social activity * Dating, a form of courtship involving social activity, with the aim of assessing a potential partner **Group dating *Play date, ...
from 80.5 million to 78.5 million years ago.Gallagher, W.B. (2005).
Recent mosasaur discoveries from New Jersey and Delaware, USA: stratigraphy, taphonomy and implications for mosasaur extinction.
" ''Netherlands Journal of Geosciences'', 84(3): 241.


References


External links


The discovery of ''H. foulkii''Hadrosaurus skeletal diagram
website
Joseph Leidy and ''Hadrodaurus foulkii'' at The Academy of Natural Sciences''Hadrosaurus foulkii'' exhibit at The Academy of Natural Sciences
{{Taxonbar, from=Q14511 Late Cretaceous dinosaurs of North America Hadrosaurs Haddonfield, New Jersey Natural history of New Jersey Fossil taxa described in 1858 Taxa named by Joseph Leidy Paleontology in New Jersey Campanian genus first appearances Campanian genus extinctions Symbols of New Jersey Ornithischian genera