Mahakala omnogovae
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''Mahakala'' ( from
संस्कृतम् Sanskrit (; attributively , ; nominally , , ) is a classical language belonging to the Indo-Aryan branch of the Indo-European languages. It arose in South Asia after its predecessor languages had diffused there from the northwest in the lat ...
(''Sanskrit''), named for
Mahakala Mahākāla is a deity common to Hinduism and Tantric Buddhism. In Buddhism, Mahākāla is regarded as the sacred '' Dharmapāla'' ("Protector of the Dharma"), while in Hinduism, Mahākāla is a fierce manifestation of the Hindu god Shiva and th ...
, one of eight protector deities (
dharmapala A ''dharmapāla'' (, , ja, 達磨波羅, 護法善神, 護法神, 諸天善神, 諸天鬼神, 諸天善神諸大眷屬) is a type of wrathful god in Buddhism. The name means "'' dharma'' protector" in Sanskrit, and the ''dharmapālas'' are a ...
s) in
Tibetan Buddhism Tibetan Buddhism (also referred to as Indo-Tibetan Buddhism, Lamaism, Lamaistic Buddhism, Himalayan Buddhism, and Northern Buddhism) is the form of Buddhism practiced in Tibet and Bhutan, where it is the dominant religion. It is also in majo ...
) 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 halszkaraptorine
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 ...
from 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 (about 80 million years ago)
Upper 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'', th ...
Djadokhta Formation The Djadochta Formation (sometimes transcribed and also known as Djadokhta, Djadokata, or Dzhadokhtskaya) is a highly fossiliferous geological formation situated in Central Asia, Gobi Desert, dating from the Late Cretaceous period, about 75 mill ...
of Ömnögovi,
Mongolia Mongolia; Mongolian script: , , ; lit. "Mongol Nation" or "State of Mongolia" () is a landlocked country in East Asia, bordered by Russia to the north and China to the south. It covers an area of , with a population of just 3.3 million, ...
. It is based on a partial skeleton found in the Gobi Desert. ''Mahakala'' was a small
dromaeosaurid Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...
, and its skeleton shows features that are also found in early
troodontids 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 ...
and
avialans 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 ...
. Despite its late appearance, it is among the most basal dromaeosaurids. Its small size, and the small size of other basal
deinonychosauria Deinonychosauria is a clade of paravian dinosaurs which lived from the Late Jurassic to the Late Cretaceous periods. Fossils have been found across the globe in North America, Europe, Africa, Asia, South America, and Antarctica,Case, J.A., Mar ...
ns, suggests that small size appeared before flight capability in
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s.


Description

''Mahakala'' was a small dromaeosaurid, measuring long and weighing . The holotype specimen,
IGM Immunoglobulin M (IgM) is one of several isotypes of antibody (also known as immunoglobulin) that are produced by vertebrates. IgM is the largest antibody, and it is the first antibody to appear in the response to initial exposure to an antig ...
100/1033, consists of a partial skeleton including skull bones,
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 characteristi ...
e, limb bones, and portions of the pelvis and
shoulder girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of ...
. Although this individual was small, comparable in size to '' Archaeopteryx'', ''
Caudipteryx ''Caudipteryx'' (which means "tail feather") is a genus of peacock-sized theropod dinosaurs that lived in the Barremian age of the early Cretaceous (about 124.6 million years ago). They were feathered and extremely birdlike in their overall appea ...
'', and '' Mei'', it was close to adulthood. This genus can be distinguished from other paravians (dromaeosaurids, troodontids, and birds) by details of 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 ...
, thighbone, ilium, and tail vertebrae. Like ''Archaeopteryx'' and derived dromaeosaurids, but unlike basal troodontids and other dromaeosaurids, the middle (third)
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 me ...
was not compressed, suggesting that the uncompressed version was the basal version. It had a typical dromaeosaurid form of the second toe, with an expanded claw.


Classification

A
phylogenetic analysis 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 o ...
performed by Turner and colleagues, who described the specimen, found ''Mahakala'' to be the most basal known dromaeosaurid. Their results, along with the small size of other
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 ...
s found at the base of paravian lineages, suggest that small size was not an innovation of early birds, but a common trait of early paravians; small size would have preceded flight and would not have been a special avian
autapomorphy 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 ...
as the result of a size squeeze. Like birds, troodontids and dromaeosaurids were not small throughout their evolutionary history, and showed size increases among several different lineages. ''Mahakala'' has a combination of characteristics found among basal troodontids and birds, but lacks some that are present in more derived dromaeosaurids. A study in 2017 found that ''Mahakala'' was a member of the enigmatic, basal subfamily
Halszkaraptorinae 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 ...
. 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 ...
below is based on the phylogenetic analysis conducted in 2017 by Cau et al. using updated data from the Theropod Working Group in their description of '' Halszkaraptor''.


Paleoecology and paleobiology

The paleoenvironment of the Djadokhta Formation is interpreted as having a
semiarid A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of semi- ...
climate, with sand dune and
alluvial Alluvium (from Latin ''alluvius'', from ''alluere'' 'to wash against') is loose clay, silt, sand, or gravel that has been deposited by running water in a stream bed, on a floodplain, in an alluvial fan or beach, or in similar settings. All ...
settings. The semiarid steppe landscape was drained by
intermittent stream Intermittent, temporary or seasonal rivers or streams cease to flow every year or at least twice every five years.(Tzoraki et al., 2007) Such rivers drain large arid and semi-arid areas, covering approximately a third of the earth's surface. ...
s and was sometimes affected by dust and
sandstorms A dust storm, also called a sandstorm, is a meteorological phenomenon common in arid and semi-arid regions. Dust storms arise when a gust front or other strong wind blows loose sand and dirt from a dry surface. Fine particles are transporte ...
, and moisture was seasonal. Animals present included terrestrial
turtle Turtles are an order of reptiles known as Testudines, characterized by a special shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) and Cryptodira (hidden necked t ...
s and
crocodilia Crocodilia (or Crocodylia, both ) is an order of mostly large, predatory, semiaquatic reptiles, known as crocodilians. They first appeared 95 million years ago in the Late Cretaceous period ( Cenomanian stage) and are the closest livi ...
ns, lizards, mammals, and a variety of dinosaurs;
aquatic animal An aquatic animal is any animal, whether invertebrate or vertebrate, that lives in water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic ani ...
s like
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of ...
were not present. The majority of the
fauna Fauna is all of the animal life present in a particular region or time. The corresponding term for plants is ''flora'', and for fungi, it is ''funga''. Flora, fauna, funga and other forms of life are collectively referred to as ''Biota (ecology ...
was small to medium-sized. Small
coelurosauria Coelurosauria (; from Greek, meaning "hollow tailed lizards") is the clade containing all theropod dinosaurs more closely related to birds than to carnosaurs. Coelurosauria is a subgroup of theropod dinosaurs that includes compsognathids, t ...
ns are the most diverse dinosaurs, including fellow dromaeosaurid ''Velociraptor'', troodontids '' Byronosaurus'' and ''
Saurornithoides ''Saurornithoides'' ( ) is a genus of troodontid maniraptoran dinosaur, which lived during the Late Cretaceous period. These creatures were predators, which could run fast on their hind legs and had excellent sight and hearing. The name is deri ...
'',
oviraptorids Oviraptoridae is a group of bird-like, herbivorous and omnivorous maniraptoran dinosaurs. Oviraptorids are characterized by their toothless, parrot-like beaks and, in some cases, elaborate crests. They were generally small, measuring between on ...
''
Citipati ''Citipati'' (; meaning "funeral pyre lord") is a genus of oviraptorid dinosaur that lived in Asia during the Late Cretaceous period, about 75 million to 71 million years ago. It is mainly known from the Ukhaa Tolgod locality at the Djadokhta F ...
'', ''
Khaan ''Khaan'' (; from Mongol 'lord') was an oviraptorid dinosaur that was found in the Djadochta Formation of Mongolia and lived in the Late Cretaceous Period (Campanian), 75-71 million years ago. Description ''Khaan'' did not differ much fro ...
'', and ''
Oviraptor ''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 Chapma ...
'', and
alvarezsaurids Alvarezsauridae is a family of small, long-legged dinosaurs. Although originally thought to represent the earliest known flightless birds, they are now thought to be an early diverging branch of maniraptoran theropods. Alvarezsaurids were highly ...
'' Kol'' and '' Shuvuuia''; other dinosaurs present included ceratopsians ''
Protoceratops ''Protoceratops'' (; ) is a genus of small protoceratopsid dinosaurs that lived in Asia during the Late Cretaceous, around 75 to 71 million years ago. The genus ''Protoceratops'' includes two species: ''P. andrewsi'' and the larger ''P. hellenik ...
'' and ''
Udanoceratops ''Udanoceratops'' (meaning "Udan's horned face") is a genus of large leptoceratopsid dinosaur that lived during the Late Cretaceous period of Mongolia, in what is now the Djadokhta Formation. Discovery ''Udanoceratops'' was first named and d ...
'', the hadrosaur ''
Plesiohadros ''Plesiohadros'' (meaning "near to hadrosaurids") is an extinct genus of hadrosauroid dinosaur. It is known from a partial skeleton including the skull collected at Alag Teg locality, from the Campanian Alagteeg Formation of southern Mongoli ...
'', and the
ankylosaurid Ankylosauridae () is a family of armored dinosaurs within Ankylosauria, and is the sister group to Nodosauridae. The oldest known Ankylosaurids date to around 122 million years ago and went extinct 66 million years ago during the Cretaceous–Pal ...
''
Pinacosaurus ''Pinacosaurus'' (meaning "Plank lizard") is a genus of ankylosaurid thyreophoran dinosaur that lived in Asia during the Late Cretaceous (Santonian-Campanian, roughly 75 million to 71 million years ago), mainly in Mongolia and China. The first r ...
''. Like other dromaeosaurids, ''Mahakala'' would have been a small active
predaceous Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
carnivore A carnivore , or meat-eater (Latin, ''caro'', genitive ''carnis'', meaning meat or "flesh" and ''vorare'' meaning "to devour"), is an animal or plant whose food and energy requirements derive from animal tissues (mainly muscle, fat and other s ...
.


See also

*
Timeline of dromaeosaurid research This timeline of dromaeosaurid research is a chronological listing of events in the history of paleontology focused on the dromaeosaurids, a group of sickle-clawed, bird-like theropod dinosaurs including animals like ''Velociraptor''. Since the ...


References


External links


HTML version of "A Basal Dromaeosaurid and Size Evolution Preceding Avian Flight"
{{Portal bar, Dinosaurs, Cretaceous Late Cretaceous dinosaurs of Asia Djadochta fauna Dromaeosaurs Fossil taxa described in 2007 Taxa named by Mark Norell