Brachylophosaurus canadensis
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''Brachylophosaurus'' ( or ; meaning "short-crested lizard",
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 ...
''brachys'' = short + ''lophos'' = crest + ''sauros'' = lizard, referring to its small crest) was a mid-sized member of the
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 ...
family of
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 ...
s. It is known from several
skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...
s and bonebed material from the
Judith River Formation The Judith River Formation is a fossil-bearing geologic formation in Montana, and is part of the Judith River Group. It dates to the Late Cretaceous, between 79 and 75.3 million years ago, corresponding to the "Judithian" land vertebrate age. It ...
of
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 Columb ...
, the
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 ...
of
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 its ...
and the
Oldman Formation The Oldman Formation is a stratigraphic unit of Late Cretaceous (Campanian stage) age that underlies much of southern Alberta, Canada. It consists primarily of sandstones that were deposited in fluvial channel and floodplain environments. It was ...
of
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest T ...
, living about 81-76.7 million years ago.


Discovery and later finds

''Brachylophosaurus'' was first named and described by
Charles Mortram Sternberg Charles Mortram Sternberg (1885–1981) was an American-Canadian fossil collector and paleontologist, son of Charles Hazelius Sternberg. Late in his career, he collected and described '' Pachyrhinosaurus'', '' Brachylophosaurus'', '' Parksosaur ...
in 1953 for a skull and partial skeleton,
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 ...
NMC 8893, which he had found in 1936 near
Steveville Steveville is a ghost town in southeastern Alberta, Canada near Brooks. In 1910, the community had a general store. Named after Steve Hall, a local homesteader, the community never attracted a large population. The Hall family operated a number of ...
in
Alberta Alberta ( ) is one of the thirteen provinces and territories of Canada. It is part of Western Canada and is one of the three prairie provinces. Alberta is bordered by British Columbia to the west, Saskatchewan to the east, the Northwest T ...
, and which was at first thought to belong to '' Gryposaurus'' (or '' Kritosaurus'' as it was known at the time). 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 specim ...
is ''Brachylophosaurus canadensis''. The generic name is derived from Greek βραχύς, ''brachys'', "short", and λόφος, ''lophos'', "crest of a helmet". The specific name refers to the provenance from Canada. Later, it was recognised that specimen FMNH PR 862, a partial skull discovered in 1922, could also be referred to ''B. canadensis''. The type specimen was uncovered in a layer of the middle
Oldman Formation The Oldman Formation is a stratigraphic unit of Late Cretaceous (Campanian stage) age that underlies much of southern Alberta, Canada. It consists primarily of sandstones that were deposited in fluvial channel and floodplain environments. It was ...
dating from about 78 million years ago. The holotype remained the only described and recognised specimen of the genus until the 1980s, when Jack Horner described a second species, ''Brachylophosaurus goodwini'', in 1988. The specific name honours preparator and collector Mark Goodwin. This species was based on a partial skull and skeleton, specimen UCMP 130139 found in the
Judith River Formation The Judith River Formation is a fossil-bearing geologic formation in Montana, and is part of the Judith River Group. It dates to the Late Cretaceous, between 79 and 75.3 million years ago, corresponding to the "Judithian" land vertebrate age. It ...
of Montana, at the Skull Crest. However, in 2005 a study by Albert Prieto-Márquez concluded that the perceived differences between the two species were either due to individual variation or the result of UCMP 130139 having been reconstructed with an upside down skull crest. ''B. goodwini'' would have been a
junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linn ...
of ''B. canadensis''. A further Canadian find was specimen TMP 1990.104.0001, a partial skeleton with skull in 1990 discovered at Milk River in Alberta and collected by Tyrrell staffer Darren Tanke and crew. ''Brachylophosaurus'' has subsequently become better known from fossils found in Montana than Alberta, however, despite its specific name ''canadensis''. These include specimens MOR 720, a braincase; MOR 794, a very complete skeleton with skull of an adult individual; and MOR 940, another braincase. Near
Malta, Montana Malta ( ) is a city in, and the county seat of, Phillips County, Montana, United States, located at the intersection of U.S. Routes 2 and 191. The population was 1,860 at the 2020 census. History After James Hill and his partners built th ...
an entire bonebed of ''Brachylophosaurus'' fossils has been uncovered containing over eight hundred specimens, that have been catalogued under number MOR 1071. In 1994 at Malta in Phillips County, amateur paleontologist Nate Murphy discovered a complete and uncrushed ''Brachylophosaurus'' skeleton which he nicknamed "Elvis". Subsequently, even more informative finds were made by Murphy and his team from the Judith River Dinosaur Institute. On 20 July 2000, specimen JRF 115H or "Leonardo", a fully articulated and partially "mummified" skeleton of a subadult ''Brachylophosaurus'', was discovered by Dan Stephenson. It is considered one of the most spectacular dinosaur finds ever, and was included in the ''Guinness Book of World Records''. They subsequently excavated "Roberta", an almost complete gracile skeleton, and "Peanut", a partially preserved juvenile with some skin impressions. "Peanut" was discovered in 2002 by Robert E. Buresh and is on display at the Institute in Malta, MT. In May 2008, Steven Cowan, public-relations coordinator at the Houston Museum of Natural Science, discovered a ''Brachylophosaurus'' skeleton subsequently dubbed "Marco" from the same area as Leonardo.


Description


Size and general build

''Brachylophosaurus'' is notable for its bony crest, which forms a horizontally flat, paddle-like plate over the top of the rear
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, th ...
."Brachylophosaurus." In: Dodson, Peter & Britt, Brooks & Carpenter, Kenneth & Forster, Catherine A. & Gillette, David D. & Norell, Mark A. & Olshevsky, George & Parrish, J. Michael & Weishampel, David B. ''The Age of Dinosaurs''. Publications International, LTD. p. 134. . Some, depending on their age, had crests that covered nearly the entire skull roof, while others had shorter, narrower crests. Some researchers suggest it was used for pushing contests, but it may not have been strong enough for this. Other notable features are a relatively small head, the unusually long lower arms and the beak of the upper jaw being wider than other hadrosaurs of that time. Apart from the above, ''Brachylophosaurus'' was a typical hadrosaur which reached an adult length of at least . In 2010, Gregory S. Paul estimated maximum length at resulting in weight of . Like other hadrosaurs, ''Brachylophosaurus'' had features like cheeks to keep fodder in the mouth and dental batteries with hundreds of stacked teeth. These teeth could be used to chew efficiently, a feature rare among reptiles, but common among some cerapodan
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 st ...
dinosaurs like ''Brachylophosaurus''.


Distinguishing traits

In 2015, Jack Horner established some distinguishing traits. Two of these are
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
, unique derived characters. The crest formed by the nasal bones is flat and paddle-shaped in adult individuals and largely or totally overhangs the supratemporal fenestrae. The rear edge of the prefrontal bone overgrowths the frontal bone and more to the rear is oriented inwards and downwards to support the base of the crest and contribute to the edge of the supratemporal fenestra. Additionally, there is one trait that is not unique in itself but forms a unique combination with the two autapomorphies: the front branch of the
lacrimal bone The lacrimal bone is a small and fragile bone of the facial skeleton; it is roughly the size of the little fingernail. It is situated at the front part of the medial wall of the orbit. It has two surfaces and four borders. Several bony landmarks of ...
is extremely elongated and with its tip only touches the
maxillary bone 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. The t ...
.Fowler, Elizabeth A. Freedman, and John R. Horner. "A New Brachylophosaurin Hadrosaur (Dinosauria: Ornithischia) with an Intermediate Nasal Crest from the Campanian Judith River Formation of Northcentral Montana." ''PLOS One'' 10.11 (2015): e0141304.
/ref>


Skeleton

The head of ''Brachylophosaurus'' is elongated. It is wide at the rear and very narrow along most of the length of the snout. The upper beak however, abruptly widens at its rear edge, forming a broad bone core for a horn sheath. The nostrils are extremely large and between them the nasal bones form a narrow tall bone wall on top of much of the snout. More to behind the nasal bones stretch out horizontally, creating a flat tongue-shaped skull crest that overgrowths and ultimately overhangs, most of the skull roof. The crest is not hollow but consists of massive bone. The crest has a low longitudinal ridge on the midline. 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 ...
, the tooth-bearing upper jaw bone, is rather elongated in front. Its tooth positions increase during the lifetime of the animal, ranging from thirty-three in younger individuals to forty-eight in 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. The teeth are stacked in a tooth battery, with up to three teeth per position. The battery forms a sharp cutting edge, bending inwards, with one or sometimes two teeth per position contributing to the attrition surface. More to behind, the lower
jugal bone 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. Ana ...
s and
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 pa ...
s flare out sidewards, so that the skull is much wider at its rear lower edges than at the top surface, resulting in a trapezium-shaped profile in posterior view.


Soft tissues

Several so-called "mummies" provide information about the soft tissues of ''Brachylophosaurus''. These "mummies" actually consist of natural casts formed in moulds in the stone matrix surrounding the skeleton, preserving the outline of the body and showing skin imprints. The best studied "mummy" has been "Leonardo", a specimen 90% of the cast surface of which is covered by imprints. Generally, the surface is close to the bones, which could be caused by desiccation before burial or the compressive action of the covering sediment. An exception is the region around the right shoulder, which shows the profile of about six centimetres thick muscles. "Leonardo" also indicates that the base of the neck was heavily muscled and that the soft tissue upper neck profile was placed in an elevated position, running much higher than was usually reconstructed in drawings which tended to follow the curvature of the vertebral column, and filling much of the bend between the front back and the head. On the snout, the remains of a broad
keratin Keratin () is one of a family of structural fibrous proteins also known as ''scleroproteins''. Alpha-keratin (α-keratin) is a type of keratin found in vertebrates. It is the key structural material making up Scale (anatomy), scales, hair, Nail ...
ous beak are visible. The skin impressions show many folds and a structure of small polygonal scales. On the back a midline frill formed by triangular or hatchet-shaped projections is present. These seem to be individually separated and are placed as extensions of each neural spine of the vertebral column. The second, third and fourth finger of the hand are contained in a shared soft tissue "mitten". Examination of the stomach of "Leonardo" also reveals that the dinosaur was parasitized by small, needle-like worms covered in fine bristles. The discovery indicates that other dinosaur species might have been hosts of similar parasites.


Classification

The following cladogram of hadrosaurid relationships was published in 2013 by
Alberto Prieto-Márquez Alberto is the Romance version of the Latinized form (''Albertus'') of Germanic ''Albert''. It is used in Italian, Portuguese and Spanish. The diminutive forms are ''Albertito'' in Spain or ''Albertico'' in some parts of Latin America, Albertin ...
et al.:


Paleobiology

In 2003, evidence of
tumor A neoplasm () is a type of abnormal and excessive growth of tissue. The process that occurs to form or produce a neoplasm is called neoplasia. The growth of a neoplasm is uncoordinated with that of the normal surrounding tissue, and persists ...
s, including
hemangioma A hemangioma or haemangioma is a usually benign vascular tumor derived from blood vessel cell types. The most common form, seen in infants, is an infantile hemangioma, known colloquially as a "strawberry mark", most commonly presenting on the ski ...
s, 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 was discovered in fossilized ''Brachylophosaurus'' skeletons. Rothschild ''et al.'' tested dinosaur vertebrae for tumors 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 ''Edmontosaurus'', ''Gilmoreosaurus'', and ''Bactrosaurus'', also tested positive. Although more than 10,000 fossils were examined in this manner, the tumors were limited to ''Brachylophosaurus'' and closely related genera. The tumors may have been caused by environmental factors or genetic propensity.


Diet

A 2008 study conducted on the famous dinosaur mummy Leonardo found that ''Brachylophosaurus'' had a diet that consisted of
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
,
conifer Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...
s,
fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes exce ...
s, and
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s like
magnolia ''Magnolia'' is a large genus of about 210 to 340The number of species in the genus ''Magnolia'' depends on the taxonomic view that one takes up. Recent molecular and morphological research shows that former genera ''Talauma'', ''Dugandiodendr ...
s. The study also found that ''Brachylophosaurus'' was a generalist herbivore; being both a browser and a
grazer Grazer may refer to: *grazer, an animal that grazes *Grazer, a native or inhabitant of Graz *GRAZER, the shoegaze band GRAZER Astronomy * Earth Grazer, Earth-grazing fireball that enters the Earth's atmosphere and leaves again * Mercury grazer, ...
, but it did more of the former rather than the latter due to the contents found in its stomach.


Paleoecology

Some of the less common hadrosaurs in the Dinosaur Park Formation of Dinosaur Provincial Park like ''Brachylophosaurus'' may represent the remains of individuals who died while migrating through the region. They might also have had a more upland habitat where they may have nested or fed.Tanke, D.H. and Brett-Surman, M.K. 2001. Evidence of Hatchling and Nestling-Size Hadrosaurs (Reptilia:Ornithischia) from Dinosaur Provincial Park (Dinosaur Park Formation: Campanian), Alberta, Canada. pp. 206-218. In: Mesozoic Vertebrate Life—New Research Inspired by the Paleontology of Philip J. Currie. Edited by D.H. Tanke and K. Carpenter. Indiana University Press: Bloomington. xviii + 577 pp.


See also

*
Timeline of hadrosaur research A timeline is a display of a list of events in chronological order. It is typically a graphic design showing a long bar labelled with dates paralleling it, and usually contemporaneous events. Timelines can use any suitable scale representin ...
Most Closely Related animals * '' Maiasaura'' * ''
Acristavus ''Acristavus'' (meaning "non-crested grandfather") is a genus of saurolophine dinosaur. Fossils have been found from the Campanian Two Medicine Formation in Montana and Wahweap Formation in Utah, United States. The type species ''A. gagslarsoni ...
'' * '' Wulagasaurus''(?) * ''
Probrachylophosaurus ''Probrachylophosaurus bergei'' is a species of large herbivorous brachylophosaurin hadrosaurid dinosaur known from the Late Cretaceous Campanian Judith River Formation, of Montana and the Foremost Formation of Alberta. The significance of this ...
''


References


External links


The Judith River Dinosaur Institute
from SkeletalDrawing.com {{Taxonbar, from=Q29057493, from2=Q731948 Late Cretaceous dinosaurs of North America Saurolophines Fossil taxa described in 1953 Taxa named by Charles Mortram Sternberg Oldman fauna Paleontology in Montana Paleontology in Alberta Campanian genus first appearances Campanian genus extinctions Ornithischian genera