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''Hesperornithoides'' (meaning "western bird form"; nicknamed "Lori") 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
troodontid Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil disco ...
theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...
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 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 Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
period.


Discovery

Although several Troodontid teeth were found, with the troodontid '' Koparion'' named in 1994, a well preserved skeleton wasn't found until in 2001, a field crew from the
Tate Museum Tate is an institution that houses, in a network of four art galleries, the United Kingdom's national collection of British art, and international modern and contemporary art. It is not a government institution, but its main sponsor is the ...
supervised by William Wahl unexpectedly discovered the type skeleton ''Hesperornithoides'' in Jimbo Quarry of the
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
, overlying the excavation site of ''
Supersaurus ''Supersaurus'' (meaning "super lizard") is a genus of diplodocid sauropod dinosaur that lived in North America during the Late Jurassic period. The type species, ''S. vivianae'', was first discovered by Vivian Jones of Delta, Colorado, in ...
vivianae'', near
Douglas, Wyoming Douglas is a city in Converse County, Wyoming, United States. The population was 6,120 at the 2010 census. It is the county seat of Converse County and the home of the Wyoming State Fair. History Douglas was platted in 1886 when the Wyoming ...
. The muddy sandstone layers of the Jimbo Quarry from which ''Hesperornithoides'' came from the middle Morrison Formation dating to the between the Oxfordian and
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by ...
ages of the
Upper Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
.Trujillo, K., Chamberlain, K., & Strickland, A. (2006). Oxfordian Park: U/Pb ages from SHRIMP analysis for the Upper Jurassic Morrison Formation of southeastern Wyoming with implications for biostratigraphic correlations. ''Journal of Vertebrate Paleontology'', ''26'', 1-152.Lovelace, D.M. (2006). "An Upper Jurassic Morrison Formation fire-induced debris flow: Taphonomy and paleoenvironment of a sauropod (Sauropoda: ''Supersaurus vivianae'') locality, east-central Wyoming." pp. 47–56 ''in'' Foster, J.R., and Lucas, S.G., eds. (2006), ''Paleontology and geology of the Upper Jurassic Morrison Formation''. New Mexico Museum of Natural History and Science Bulletin 36. The accidental discovery of the skeleton led to some of the fossils being damaged or lost. 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 ...
, WYDICE-DML-001 consists of a semi-articulated partial skeleton of a subadult or adult including: the rear skull and lower jaws,
hyoid bone The hyoid bone (lingual bone or tongue-bone) () is a horseshoe-shaped bone situated in the anterior midline of the neck between the chin and the thyroid cartilage. At rest, it lies between the base of the mandible and the third cervical verteb ...
s, five cervical vertebrae, the first dorsal vertebra, twelve caudal vertebrae, a rib, chevrons, the left shoulder girdle, the right humerus, the left arm, a femur fragment, and the left and right lower legs minus the right toes. Its discovery was announced at the 2003 annual meeting of the
Society of Vertebrate Paleontology The Society of Vertebrate Paleontology (SVP) is a professional organization that was founded in the United States in 1940 to advance the science of vertebrate paleontology around the world. Mission and Activities SVP has about 2,300 members inte ...
and in 2005, a
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...
analysis including it was presented in an abstract for the ''
Journal of Vertebrate Paleontology The ''Journal of Vertebrate Paleontology'' is a bimonthly peer-reviewed scientific journal that was established in 1980 by Jiri Zidek (University of Oklahoma). It covers all aspects of vertebrate paleontology, including vertebrate origins, evolu ...
''. The same year the specimen was donated to the Big Horn Basin Foundation which in 2016 joined the
Wyoming Dinosaur Center The Wyoming Dinosaur Center is located in Thermopolis, Wyoming and is one of the few dinosaur museums in the world to have excavation sites within driving distance. The museum displays the Thermopolis Specimen of ''Archaeopteryx'', which is the ...
. In 2019, 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 ...
''Hesperornithoides miessleri'' was named and described by Scott Hartman, Mickey Mortimer, William Wahl, Dean R. Lomax, Jessica Lippincott and David M. Lovelace in an article published on ''
PeerJ ''PeerJ'' is an open access peer-reviewed scientific mega journal covering research in the biological and medical sciences. It is published by a company of the same name that was co-founded by CEO Jason Hoyt (formerly at Mendeley) and publisher ...
''. The generic name is derived from the Greek Ἑσπερίς, Hesperis, "western", ὄρνις, ''ornis'', "bird", and ~eides, "form". The specific name honours the Miessler family for its support of the project.


Description

The length of the holotype individual was in 2019 estimated at long. Such a body size is rather limited for a troodontid. The specimen may represent an adult or subadult individual, due to the lack of cranial and body proportions associated with juvenile individuals. Nevertheless, either assignation would not be much different; for instance, the estimated would become . The describing authors established some derived or
apomorphic In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
traits relative to the
Paraves Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the avial ...
. The
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 ...
is pneumatised. The rear 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 short, with less than 15% of the length of the descending branch, measured from the inner corner downwards. 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 pa ...
is part of the outer edge of the ''foramen paraquadraticum''. In the lower jaw the external side opening is small, with less than an eighth of the length of the jaw as a whole. On 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 r ...
, the ridge towards the inner condyle represents more than 15% of the total distal width of the element. The third claw of the hand is about as large as the second claw. The lower inner corner of the shinbone projects to the front.


Classification

The phylogenetic analysis of 2005 placed the specimen as a close relative of ''
Sinornithoides ''Sinornithoides'' (meaning "Chinese bird form") is a genus of troodontid theropod dinosaurs containing the single species ''Sinornithoides youngi''. ''S. youngi'' lived during the Early Cretaceous (Aptian/Albian stage, around 113 million years ...
''.Hartman, S., Lovelace, D., and Wahl, W., (2005). "Phylogenetic assessment of a maniraptoran from the Morrison Formation." ''Journal of Vertebrate Paleontology'', 25, Supplement to No. 3, pp. 67A–68A http://www.bhbfonline.org/AboutUs/Lori.pdf Unpublished phylogenetic results presented at the SVP 2017 conference recovered ''Hesperornithoides'' (then known by the informal name "Lori") as a relative of ''
Sinovenator ''Sinovenator'' (meaning "Chinese hunter") is a genus of troodontid dinosaur from China. It is from the early Cretaceous Period. Discovery and naming Two specimens of a troodontid were described in 2002. They are both housed in the Institu ...
''. The presence of this derived
maniraptoran 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 ...
along with several others, such as ''
Anchiornis ''Anchiornis'' is a genus of small, four-winged paravian dinosaurs, with only one known species, the type species ''Anchiornis huxleyi'', named for its similarity to modern birds. The Latin name ''Anchiornis'' derives from a Greek word meaning " ...
'' and ''
Eosinopteryx ''Eosinopteryx'' is an extinct genus of theropod dinosaurs known to the late Jurassic period of China. It contains a single species, ''Eosinopteryx brevipenna''. Discovery and naming ''E. brevipenna'' is known from a single fossil specimen recov ...
'', in Jurassic sediments was a strong refutation of the
temporal paradox A temporal paradox, time paradox, or time travel paradox is a paradox, an apparent contradiction, or logical contradiction associated with the idea of time and time travel. The notion of time travel to the future complies with current understanding ...
argument used by those who oppose the consensus view that birds evolved from dinosaurs. In 2019, ''Hesperornithoides'' was placed in the
Troodontidae Troodontidae is a clade of bird-like theropod dinosaurs. During most of the 20th century, troodontid fossils were few and incomplete and they have therefore been allied, at various times, with many dinosaurian lineages. More recent fossil discov ...
, in a relatively basal position, in a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
with ''
Daliansaurus ''Daliansaurus'' (meaning " Dalian reptile") is a genus of small troodontid theropod dinosaur, measuring approximately long, from the Early Cretaceous of China. It contains a single species, ''D. liaoningensis'', named in 2017 by Shen and c ...
'', ''
Xixiasaurus ''Xixiasaurus'' () is a genus of troodontid dinosaur that lived during the Late Cretaceous Period in what is now China. The only known specimen was discovered in Xixia County, Henan Province, in central China, and became the holotype of th ...
'' and '' Sinusonasus'', as shown in the cladogram below:


Palaeobiology

Many large dinosaurs have been discovered in the Morrison Formation during the nineteenth century; ''Hesperornithoides'' proves that the full wealth of species had not yet been revealed. It lived in a relatively open well-watered landscape with low growths but few trees, or,
lacustrine A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much larger ...
and
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (Anoxic waters, anoxic) processes prevailing, especially in t ...
environments. The layers of the Jimbo Quarry have been interpreted as a marginal lacustrine or
wetland A wetland is a distinct ecosystem that is flooded or saturated by water, either permanently (for years or decades) or seasonally (for weeks or months). Flooding results in oxygen-free (Anoxic waters, anoxic) processes prevailing, especially in t ...
environment, representing a cyclical rise and fall in water layers. Notably, the specimen preserves some keratin sheaths on the distal end of several pedal and manual unguals. The relative positioning of the thoracic elements of the skeleton suggests the animal was in a resting pose and the wetland environment from which the fossil was found suggests a wetland paleoecology. Apart from the
Avialae Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to deinonychosaurs, though alternative definitions are occasionally use ...
, most paravian groups had only few members with some flight capability. The exceptions all showed adaptations for an arboreal lifestyle, indicating flight developed multiple times when species evolved traits and behaviours useful for climbing trees or other more vertical surfaces, such as wing-assisted incline running.


See also

*
Timeline of troodontid research This timeline of troodontid research is a chronological listing of events in the history of paleontology focused on the troodontids, a group of bird-like theropod dinosaurs including animals like ''Troodon''. Troodontid remains were among the fir ...
*
Morrison Formation The Morrison Formation is a distinctive sequence of Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandstone, siltston ...
*
2019 in paleontology Flora Plants Fungi Paleomycological research * Fossil sporocarps indistinguishable from sporocarps of members of the extant genus ''Stemonitis'' are described from the Cretaceous amber from Myanmar by Rikkinen, Grimaldi & Schmidt (2019). * ...


References

{{Taxonbar, from=Q3301304 Late Jurassic dinosaurs of North America Troodontids Dinosaurs of the Morrison Formation Paleontology in Wyoming Fossil taxa described in 2019