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''Falcarius'' (meaning "sickle cutter") 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 ...
of primitive
therizinosaur Therizinosaurs (once called segnosaurs) were large herbivorous Theropoda, theropod dinosaurs whose fossils have been found across the Early Cretaceous, Early to Late Cretaceous deposits in Asia and North America. Various features of the forelimbs ...
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 ...
that lived during the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
period in what is now
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 the Car ...
. Its remains were first collected in the
Cedar Mountain Formation The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid-Cretaceous. The formation was named for Cedar Mountain (Utah), Cedar Mountain in northern Emery Coun ...
in 1999, with subsequent findings made during the 2000s. The genus is known from multiple specimens ranging from immature to fully-grown individuals. ''Falcarius'' was a long bipedal
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
with a small head and an elongated neck and tail. Unlike advanced therizinosaurs, ''Falcarius'' had a propubic pelvis and three-toed feet with a reduced
hallux Toes are the digits (fingers) of the foot of a tetrapod. Animal species such as cats that walk on their toes are described as being '' digitigrade''. Humans, and other animals that walk on the soles of their feet, are described as being '' pl ...
(first digit). ''Falcarius'' is the basalmost known definitive therizinosaurian genus, and has been considered a
transitional form A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
connecting the typical theropod bodyplan to the unusual morphology of
Therizinosauridae Therizinosauridae (meaning 'scythe lizards')Translated paper
is a family of derived (advanc ...
. Its description in 2005, following that of the basal
therizinosauroid Therizinosaurs (once called segnosaurs) were large herbivorous theropod dinosaurs whose fossils have been found across the Early to Late Cretaceous deposits in Asia and North America. Various features of the forelimbs, skull and pelvis unite thes ...
''
Beipiaosaurus ''Beipiaosaurus'' is a genus of therizinosauroid theropod dinosaurs that lived in Asia during the Early Cretaceous in the Yixian Formation. The first remains were found in 1996 and formally described in 1999. Before the discovery of ''Yutyrannus ...
'' from the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
of
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
in 1999, helped clarify the early evolution of the Therizinosauria and its relationships within the larger group of Theropoda.


History of discovery

The remains of ''Falcarius'' were first discovered in 1999 by commercial fossil collector Lawrence Walker at the Crystal Geyser Quarry site in Grand County. He informed paleontologist James I. Kirkland of the find, who with a team of the
Utah Geological Survey The Utah Geological Survey is based in Salt Lake City, Utah, United States. It also has an office in Cedar City, Utah. It is a division of the
from 2001 onwards began to uncover the bones that proved to be present in a two-
acre The acre is a unit of land area used in the imperial Imperial is that which relates to an empire, emperor, or imperialism. Imperial or The Imperial may also refer to: Places United States * Imperial, California * Imperial, Missouri * Imp ...
(8,000-square-meter) area of Utah's
Cedar Mountain Formation The Cedar Mountain Formation is the name given to a distinctive sedimentary geologic formation in eastern Utah, spanning most of the early and mid-Cretaceous. The formation was named for Cedar Mountain (Utah), Cedar Mountain in northern Emery Coun ...
(Yellow Cat member). ''Falcarius'' thus lived in the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
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 ...
. Two extensive bone beds were discovered, including the remains of hundreds, perhaps thousands, of individuals of the new species. The minimum number of individual animals was in 2006 estimated at three hundred. In 2005 over two thousand specimens had been excavated, mostly consisting of disarticulated bones. These included the remains of juvenile animals. In 2008 a second site was reported, the Suarez Quarry, with mainly adult individuals, but of a perhaps slightly different type. In 2010 the number of specimens from the original quarry had increased to over 2,700, and was later that year reported to have risen to over three thousand. The first remains were partially described and discussed in 2004 on several abstracts by
paleontologists 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 ...
Lindsay E. Zanno Lindsay E. Zanno is an American vertebrate paleontology, paleontologist and a leading expert on Theropoda, theropod dinosaurs and Cretaceous paleoecosystems. She is the Head of Paleontology at the North Carolina Museum of Natural Sciences and an ...
, then a doctoral student at the university,
James I. Kirkland James Ian Kirkland (born August 24, 1954) is an American Palaeontologist, paleontologist and geologist. He has worked with dinosaur remains from the south west United States of America and Mexico and has been responsible for discovering new and i ...
,
Scott D. Sampson Scott Donald Sampson (born April 22, 1961) is a Canadian paleontologist and science communicator. Sampson is currently the Executive Director of California Academy of Sciences in San Francisco, California. He was previously Vice President of Res ...
, chief curator at the
University of Utah The University of Utah (U of U, UofU, or simply The U) is a public research university in Salt Lake City, Utah. It is the flagship institution of the Utah System of Higher Education. The university was established in 1850 as the University of De ...
's
Utah Museum of Natural History The Natural History Museum of Utah (NHMU) is a museum located in Salt Lake City, Utah, United States. The museum shows exhibits of natural history subjects, with an emphasis on Utah and the Intermountain West. The mission of the museum is to il ...
, Donald D. DeBlieux, David K. Smith and R. Kent Sanders. It was not formally named until a subsequent paper published in the May 2005 issue of the journal ''
Nature Nature, in the broadest sense, is the physics, physical world or universe. "Nature" can refer to the phenomenon, phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. ...
''. Zanno subsequently and informally described ''Falcarius'' as "the ultimate in bizarre: a cross between an ostrich, a gorilla, and Edward Scissorhands." The generic name, ''Falcarius'', is literally taken from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
falcarius (meaning sickle cutter) in reference to the large hand claws. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, ''utahensis'', reflects the provenance from Utah. 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, UMNH VP 15000, consists of a partial braincase and a large number of
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
s were assigned. ''Falcarius'' has since been described in detail by Zanno in a series of studies, beginning with her thesis in 2004.


Description

''Falcarius'' is known from multiple specimens at different ontogenic stages. While the smallest specimen was approximately less than , average adults were about in length weighing approximately .


Skull

The head anatomy of ''Falcarius'' is partially known. The skull was small and elongated. With its long neck, ''Falcarius'' could apparently reach about off the ground to munch leaves or fruit. The teeth numbered at least sixteen in 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. The t ...
of the upper jaw. 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 ...
(part of the lower jaw) carried twenty-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, tear ...
. Its small leaf-shaped and very finely serrated maxillary teeth indicate that it consumed plant material. The front-most five teeth of the lower jaw are much longer, straighter and more pointed though, and might indicate a partially omnivorous diet including meat, e.g. small animals such as
lizard Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia alt ...
s. In the back of the head, the braincase was relatively large. Its lower elements were moderately inflated by pneumatised, hollowed-out, bone tissue. The large skull opening in the snout, the ''fenestra antorbitalis'', is positioned in a depression which reaches the side of the
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
. There are at least five pairs of conical teeth in the front of the lower jaws. These teeth are hollowed-out at the inside. On the underside of the vertebrae of the neck there is a depression with at the mid-line a ridge. On the underside of the braincase there is a well-developed depression, the ''recessus basisphenoidalis''. At the back of the head there are deep depressions below the
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
and on the subotics, each with several pneumatic grooves.


Postcranial skeleton

The partitioning of the vertebral column followed that of most maniraptorans. The neck was very long with elongated cervical vertebrae. The tail was relatively long. The arm was moderately long with a somewhat robust
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 roun ...
. The relatively large, and slightly recurved, pointed ten- to thirteen-centimeter (four- to five-inch) hand claws were likely used in self-defence. ''Falcarius'' is known from many specimens, including complete forelimb specimens. Most of the bones of the pectoral girdle and forelimb are known, although sternal bones are not preserved. Both a left and right scapula are preserved, and they are both mostly complete as well. The blades of the scapulae are about long, and also quite slender, with barely any expansion. They are curved instead of straight. The scapula is placed in a relatively high position. Two coracoids are also preserved, although they differ more than the scapulae in morphology. The right coracoid is the better preserved of the two. The coracoid is not specially enlarged. The
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
is elongated and narrow, its branches placed at an angle of 104°. It is V-shaped and possesses a small hypocleidium at the front. The arm as a whole is not especially developed in length but shows signs of an increasing robustness. The humerus has a rather short triangular deltopectoral crest. The epicondyles are exceptionally thick, indicating a strong musculature. The
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 ...
is lightly curved. The hand is rather long but not very robust, with in general an elongated
metacarpus In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ...
, although the first
metacarpal In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpal bones ...
is short. The first finger diverges from the second finger, the longest of the hand. The third finger is very thin. The hand claws are moderately curved, rather pointed, and moderately long. The vertebrae of the back have below their front extensions, the
prezygapophyses The articular processes or zygapophyses (Greek language, Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "-physis, process") of a vertebra are projections of the vertebra that serve the purpose of fitting ...
, a groove which is divided in three smaller depressions. On the thumb claw the raised attachment point for the tendon of the extensor muscle is bordered by deep grooves for
ligament A ligament is the fibrous connective tissue that connects bones to other bones. It is also known as ''articular ligament'', ''articular larua'', ''fibrous ligament'', or ''true ligament''. Other ligaments in the body include the: * Peritoneal li ...
s. The middle vertebrae of the back have a second ridge extending from the base of the upper rib joint, the diapophysis, to the back of the vertebra. In the arm, the shaft of the
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 roun ...
is reinforced by powerful, somewhat obliquely placed, thickenings extending towards the
condyle A condyle (;Entry "condyle"
in
pubis. In the pelvis the ilium was by a moderate inclination partly adapted to a more raised position of the trunk. The pelvis was propubic or mesopubic: the
pubic bone In vertebrates, the pubic region ( la, pubis) is the most forward-facing (ventral and anterior) of the three main regions making up the coxal bone. The left and right pubic regions are each made up of three sections, a superior ramus, inferior ra ...
pointed more or less forwards. The leg was relatively long and adapted to running, with the lower leg longer than the thighbone. In the foot there were three weight-bearing toes. The first toe did not reach the ground nor did the first
metatarsal 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 med ...
touch the ankle.


Classification

The describers in 2005 assigned ''Falcarius'' to the
Therizinosauroidea Therizinosaurs (once called segnosaurs) were large herbivorous theropod dinosaurs whose fossils have been found across the Early to Late Cretaceous deposits in Asia and North America. Various features of the forelimbs, skull and pelvis unite thes ...
''
sensu ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...
'' of Zhang, in a basal position. Zanno later placed it as the basalmost taxon in the Therizinosauria, just outside of Therizinosauroidea ''sensu'' of Sereno. Because of the primitive position and relative completeness of ''Falcarius'', it is a good taxon to use to compare to related groups and descendant taxa. ''Falcarius'' resembles dinosaurs belonging to the
Therizinosauridae Therizinosauridae (meaning 'scythe lizards')Translated paper
is a family of derived (advanc ...
, part of the group
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 ...
. ''Falcarius'' itself probably does not belong to this former group, although it does belong to the more inclusive group
Therizinosauria Therizinosaurs (once called segnosaurs) were large herbivorous theropod dinosaurs whose fossils have been found across the Early to Late Cretaceous deposits in Asia and North America. Various features of the forelimbs, skull and pelvis unite the ...
. The group is characterized by wider hips, a comparatively large
braincase In human anatomy, the neurocranium, also known as the braincase, brainpan, or brain-pan is the upper and back part of the skull, which forms a protective case around the brain. In the human skull, the neurocranium includes the calvaria or skul ...
and long necks typical of plant eaters. The less basal Asian specimens were covered in quill-like
feather Feathers are epidermal growths that form a distinctive outer covering, or plumage, on both avian (bird) and some non-avian dinosaurs and other archosaurs. They are the most complex integumentary structures found in vertebrates and a premier ...
s; this is assumed for ''Falcarius'' as well. In general ''Falcarius'' can be seen as a
transitional form A transitional fossil is any fossilized remains of a life form that exhibits traits common to both an ancestral group and its derived descendant group. This is especially important where the descendant group is sharply differentiated by gross a ...
between the original theropod build and the more derived
Therizinosauridae Therizinosauridae (meaning 'scythe lizards')Translated paper
is a family of derived (advanc ...
. ''Falcarius'' shared some derived traits with the therizinosaurids: it had a long neck, small head with teeth adapted for eating plant material, a more robust arm with large hand-claws and a more vertical position of the body. On the other hand, it resembled more typical theropods in having a long tail, a propubic pelvis, a long lower leg and a foot without a fourth toe reaching the ground. The
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 d ...
below shows the results of a phylogenetic analysis of Therizinosauria performed by Yoshitsugu Kobayashi and team in 2022, based on their description of ''
Paralitherizinosaurus ''Paralitherizinosaurus'' is an extinct genus of therizinosaurid dinosaur from the Late Cretaceous Osoushinai Formation of Hokkaido, Japan. The genus contains a single species, ''P. japonicus'', known from a partial right hand and cervical vertebr ...
''. ''Falcarius'' is recovered as the basalmost therizinosaurian taxon:


See also

*
Timeline of therizinosaur research The timeline of therizinosaur research is a chronological listing of events in the history of paleontology focused on therizinosaurs. They were unusually long-necked, pot-bellied, and large-clawed herbivorous theropods most closely related to bir ...


References


External links


BBC


{{Taxonbar, from=Q251225 Early Cretaceous dinosaurs of North America Fossil taxa described in 2005 Therizinosaurs Transitional fossils Taxa named by James I. Kirkland Taxa named by James M. Clark Taxa named by Scott D. Sampson Paleontology in Utah Taxa named by Lindsay Zanno