Halszkaraptorines Temporal range: Late Cretaceous, | |
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Holotype specimen of Halszkaraptor escuilliei | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Family: | † Dromaeosauridae |
Subfamily: | † Halszkaraptorinae Cau et al., 2017 |
Type genus | |
† Halszkaraptor escuilliei Cau et al., 2017 | |
Genera | |
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Halszkaraptorinae is a basal ("primitive") subfamily of Dromaeosauridae (or possibly Unenlagiidae) that includes the enigmatic genera Halszkaraptor , Natovenator , Mahakala , and Hulsanpes . Halszkaraptorines are definitively known only from Late Cretaceous strata in Asia, specifically in Mongolia. It is debated among researchers whether the group had a semiaquatic lifestyle.
The first known remains of halszkaraptorines were found in sandstone sediments in 1970 during a Polish-Mongolian expedition at the Barun Goyot Formation, Gobi Desert. Later in 1982, they were described by Polish palaeontologist Halszka Osmólska and used as the holotype for the new genus and species Hulsanpes perlei , honoring the Mongolian paleontologist Altangerel Perle. Although the affinities of these fossils were not fully understood, they were tentatively interpreted as belonging to Deinonychosauria. [1] In 1992 dromaeosaurid-like fossils were recovered from the Tugriken Shire locality of the Djadokhta Formation and described as a new species in 2007 by Alan H Turner and team. This new taxon, Mahakala omnogovae , was regarded as a primitive dromaeosaurid mainly based on its body proportions. [2] In 2015 the Royal Belgian Institute of Natural Sciences received a small theropod fossil specimen, hailing from England and Japanese private collections. The specimen, through negotiations between the former institute, Eldonia and Mongolian authorities, was lastly returned to the Institute of Paleontology and Geology, Mongolian Academy of Sciences. According to associated documents of the specimen, it was illegally poached from the Djadokhta Formation (Ukhaa Tolgod locality; however, sediments may suggest that, alternatively, it was recovered from nearby Bayn Dzak locality) during an unknown year, eventually falling into private collections. The specimen was formally described in 2017 by Andrea Cau with colleagues, naming the genus and species Halszkaraptor escuilliei , in honor of Halszka Osmólska and François Escuillié, paleontologists who made negotiations for returning the poached specimen to Mongolia possible. With the naming of Halszkaraptor, the subfamily Halszkaraptorinae was also coined to contain this taxon and relatives, namely Hulsanpes and Mahakala. [3]
Halszkaraptorines were relatively small theropods, reaching lengths similar to those of modern-day ducks. [2] [3] They are diagnosed by their long necks, proximal caudal vertebrae with oriented articular processes (projections of the vertebra fits with an adjacent vertebra) and prominent zygapophyseal laminae (plates of bone that form the posterior walls of each vertebra), a flattened ulna with a sharp posterior margin, ilium with a shelf-like supratrochanteric (above the trochanter of the femur) process, metacarpal III shaft transversely as thick as that of metacarpal II, a posterodistal surface on the femoral shaft with an elongate fossa bound by a lateral crest and the proximal half of metatarsal III being unconstricted and markedly convex anteriorly. [3]
Halszkaraptorinae is defined as the most inclusive clade that contains Halszkaraptor escuilliei but not Dromaeosaurus albertensis, Unenlagia comahuensis, Saurornithoides mongoliensis or Vultur gryphus . [3] The cladogram below is based on the phylogenetic analysis conducted in 2022 by Lee et al. in their description of Natovenator . [4]
In 2017 a comparison of the fossils of Halszkaraptor with the bones of extant crocodilians and aquatic birds revealed evidence of a semiaquatic lifestyle. Cau and colleagues suggested that halszkaraptorines were obligate bipeds on land, but also swimmers that used its forelimbs to push through water and used their long necks for foraging. Such traits allowed them to exploit both terrestrial and aquatic paleoecosystems. [3] A 2024 study by Tse, Miller, and Pittman, focusing on the skull morphology and bite forces of various dromaeosaurids discovered that Halszkaraptor had a rapid bite unsuited to piscivorous feeding as previously hypothesized based on its skull morphology, and instead suggest it was an insectivore that hunted small invertebrates possibly in low-light conditions (at night or in murky water), since it likely had exceptional low-light vision among dromaeosaurids based on its relatively large orbit size. [5]
Other researchers have either disagreed with or merely followed Cau's interpretation as a semiaquatic dinosaur. In 2019, Brownstein argued that the features noted for Halszkaraptor do not directly support its ability to swim. He also suggested that this dinosaur may be a basal dromaeosaur with transitional features, [6] although Cau rebutted his claims a year later. [7] In 2021, Hone and Holtz noted that since Halszkaraptor and many modern aquatic birds with no flattened unguals is said to be semi-aquatic, having flattened unguals like Spinosaurus do not necessarily suggest that the animal can swim; they didn't propose their own view on this dinosaur's potential ability to swim. [8] In 2022, Fabbri and his colleagues argued against a semi-aquatic ecology for Halszkaraptor, noting that it had low bone density, a trait not observed in semi-aquatic animals. [9] In response, Cau has pointed out on his blog that swans similarly have low bone density yet have adaptations for semi-aquatic feeding. [10]
Velociraptor is a genus of small dromaeosaurid dinosaurs that lived in Asia during the Late Cretaceous epoch, about 75 million to 71 million years ago. Two species are currently recognized, although others have been assigned in the past. The type species is V. mongoliensis, named and described in 1924. Fossils of this species have been discovered in the Djadochta Formation, Mongolia. A second species, V. osmolskae, was named in 2008 for skull material from the Bayan Mandahu Formation, China.
Troodontidae is a clade of bird-like theropod dinosaurs from the Late Jurassic to Late Cretaceous. 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 discoveries of complete and articulated specimens, have helped to increase understanding about this group. Anatomical studies, particularly studies of the most primitive troodontids, like Sinovenator, demonstrate striking anatomical similarities with Archaeopteryx and primitive dromaeosaurids, and demonstrate that they are relatives comprising a clade called Paraves.
Oviraptor is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous period. The first remains were collected from the Djadokhta Formation of Mongolia in 1923 during a paleontological expedition led by Roy Chapman Andrews, and in the following year the genus and type species Oviraptor philoceratops were named by Henry Fairfield Osborn. The genus name refers to the initial thought of egg-stealing habits, and the specific name was intended to reinforce this view indicating a preference over ceratopsian eggs. Despite the fact that numerous specimens have been referred to the genus, Oviraptor is only known from a single partial skeleton regarded as the holotype, as well as a nest of about fifteen eggs and several small fragments from a juvenile.
Avimimus, meaning "bird mimic", is a genus of oviraptorosaurian theropod dinosaur, named for its bird-like characteristics, that lived in the late Cretaceous in what is now Mongolia, around 85 to 70 million years ago.
Bagaceratops is a genus of small protoceratopsid dinosaurs that lived in Asia during the Late Cretaceous, around 72 to 71 million years ago. Bagaceratops remains have been reported from the Barun Goyot Formation and Bayan Mandahu Formation. One specimen may argue the possible presence of Bagaceratops in the Djadochta Formation.
Borogovia is a troodontid theropod dinosaur genus which lived during the Late Cretaceous Period, in what is now Mongolia.
Tylocephale is a genus of pachycephalosaurid dinosaur, a group of dome-headed, herbivorous ornithischians, that lived during the Late Campanian stage of the Late Cretaceous in what is now Mongolia. It is known from a partial skull and associated mandible that were unearthed in 1971 by a Polish-Mongolian Expedition to the Barun Goyot Formation of the Gobi Desert. The specimen was described in 1974 by Polish paleontologists Teresa Maryańska and Halszka Osmólska as a new genus and species.
Kuru is a genus of dromaeosaurid theropod from the Late Cretaceous Barun Goyot Formation of Mongolia. The genus contains only a single species, the type species Kuru kulla, which is known from a fragmentary skeleton including a partial skull.
Heyuannia is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous epoch, in what is now China and Mongolia. It was the first oviraptorid found in China; most others were found in neighbouring Mongolia. Two species are known: H. huangi, named by Lü Junchang in 2002 from the Dalangshan Formation; and H. yanshini, originally named as a separate genus Ingenia from the Barun Goyot Formation by Rinchen Barsbold in 1981, and later renamed to Ajancingenia in 2013 due to the preoccupation of Ingenia. The latter name was eventually discarded due to various ethical issues surrounding the author.
Hulsanpes is a genus of halszkaraptorine theropod dinosaurs that lived during the Late Cretaceous in what is now the Barun Goyot Formation of Mongolia, about 75-72 million years ago. The remains were found in 1970 and formally described in 1982 by Halszka Osmólska, who noted that the genus is represented by an immature individual. Hulsanpes represents the first record of the basal dromaeosaurid subfamily Halszkaraptorinae.
The Barun Goyot Formation is a geological formation dating to the Late Cretaceous Period. It is located within and is widely represented in the Gobi Desert Basin, in the Ömnögovi Province of Mongolia.
The Djadochta Formation is a highly fossiliferous geological formation situated in Central Asia, Gobi Desert, dating from the Late Cretaceous period, about 75 million to 71 million years ago. The type locality is the Bayn Dzak locality, famously known as the Flaming Cliffs. Reptile and mammal remains are among the fossils recovered from the formation.
Halszka Osmólska was a Polish paleontologist who had specialized in Mongolian dinosaurs.
Mahakala is a genus of halszkaraptorine theropod dinosaur from the Campanian-age Upper Cretaceous Djadokhta Formation of Ömnögovi, Mongolia. It is based on a partial skeleton found in the Gobi Desert. Mahakala was a small dromaeosaurid, and its skeleton shows features that are also found in early troodontids and avialans. Despite its late appearance, it is among the most basal dromaeosaurids. Its small size, and the small size of other basal deinonychosaurians, suggests that small size appeared before flight capability in birds. The genus is named for Mahakala, one of eight protector deities (dharmapalas) in Tibetan Buddhism.
Ceratonykus is a monospecific genus of alvarezsaurid dinosaur from Mongolia that lived during the Late Cretaceous in what is now the Barun Goyot Formation. The type and only species, Ceratonykus oculatus, is known from a fragmentary skeleton, including an incomplete skull, of an adult individual. It was named and described in 2009 by Vladimir Alifanov and Rinchen Barsbold. Its describers questioned the traditional placement of alvarezsaurs in Theropoda, instead suggesting they were ornithischians, but this has not been accepted since. Ceratonykus has an estimated length of 75 centimetres and weight of 760 grams. It has been considered as a possible junior synonym of Parvicursor.
Unenlagiidae is a proposed family of eumaniraptoran paravians that includes the subfamilies Unenlagiinae and possibly Halszkaraptorinae. Fossils of both subfamilies have been found in both Gondwanan and Laurasian deposits. The biology of the group suggests that some members were semiaquatic specialists.
Halszkaraptor is a genus of waterfowl-like dromaeosaurid dinosaurs from Mongolia that lived during the Late Cretaceous period. It contains only one known species, Halszkaraptor escuilliei.
Ondogurvel is a genus of alvarezsaurid dinosaur from the Late Cretaceous (Campanian) Barun Goyot Formation in southern Mongolia. The type and only species is O. alifanovi, known from a partial skeleton consisting of fragments of two last dorsal vertebrae, three anterior sacral vertebrae, right ilium, left and right pubis and ischium, articulated right tibia, fibula, metatarsals II and IV, and phalanges IV-1 and IV-2, right carpometacarpus, left and right manual phalanx II-1, right femur, left pedal phalanx II-1, and fragments of unidentified phalanges.
Natovenator is a genus of halszkaraptorine dromaeosaurid dinosaur from the Late Cretaceous Barun Goyot Formation of Mongolia. The genus is known from a single species, N. polydontus. Natovenator is crucial to the understanding of halszkaraptorines due to it providing more support for the semi-aquatic lifestyle that has been proposed for this clade. This discovery is important as the semi-aquatic lifestyles of halszkaraptorines was contested in early 2022.
Jaculinykus is an extinct genus of alvarezsaurid theropod dinosaur from the Late Cretaceous Baruungoyot Formation of Mongolia. The genus contains a single species, J. yaruui, known from a nearly complete articulated skeleton including bones of the skull. Jaculinykus is notable for its unique hand, which has a hypertrophied first digit and greatly reduced second digit, which is intermediate between the tridactyl hand of Shuvuuia and monodactyl hand of Linhenykus.