megaraptorid Megaraptora is a clade of carnivorous tetanuran theropod dinosaurs with controversial relations to other theropods. Its derived members, the Megaraptoridae are noted for their elongated hand claws and proportionally large arms, which are usually ...

Late 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'', the ...

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...

Chorrillo Formation The Chorrillo Formation, also named as Chorillo Formation,Río Leona
at Santa Cruz,

Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...

. The genus contains a single species, ''M. macrothorax'', known from an incomplete, disarticulated skeleton. ''Maip'' may represent the largest megaraptorid known from

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

, and possibly the world.

Discovery and naming

The ''Maip''

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, MPM 21545, was discovered by Alexis Rolando on the La Anita Farm, west of El Calafate, Santa Cruz province, Argentina, in 2019. The specimen was found disarticulated but in association over an area of . The known fossil material consists of the axis, several dorsal and

caudal Caudal may refer to: Anatomy * Caudal (anatomical term) (from Latin ''cauda''; tail), used to describe how close something is to the trailing end of an organism * Caudal artery, the portion of the dorsal aorta of a vertebrate that passes into the ...

vertebrae, cervical and dorsal ribs, gastralia, a left coracoid, fragmentary

scapula The scapula (plural scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on eithe ...

, partial right pubis, and partial metatarsal. Some of these bones were described in 2019 by Novas ''et al''. The holotype specimen includes some bones previously unknown in other megaraptorids. Although fragmentary, it represents one of the most complete megaraptorid skeletons known. ''Maip'' was first announced in a Research Square

preprint In academic publishing, a preprint is a version of a scholarly or scientific paper that precedes formal peer review and publication in a peer-reviewed scholarly or scientific journal. The preprint may be available, often as a non-typeset versio ...

in 2021. However, since the paper did not meet the necessary

requirements In product development and process optimization, a requirement is a singular documented physical or functional need that a particular design, product or process aims to satisfy. It is commonly used in a formal sense in engineering design, includi ...

, the taxon was considered informally named. In 2022, the fossil material was validly described as belonging to a new genus and species of megaraptorid by Rolando ''et al''. The generic name, "''Maip''", references a malicious being in

Aonikenk Tehuelche (''Aoniken, Inaquen, Gunua-Kena, Gununa-Kena'') is one of the Chonan languages of Patagonia. Its speakers were nomadic hunters who occupied territory in present-day Chile, north of Tierra del Fuego and south of the Mapuche people. It is ...

mythology that is “the shadow of death” that “kills with cold wind." The specific name, "''macrothorax''", is derived from the Greek "makrós", meaning "long", and 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 ...

"thorax", meaning "chest", in reference to its large thoracic cavity.

Description

''Maip'' is estimated to have been around long in life. As such, it may be the largest megaraptorid currently known. Members of the Megaraptoridae as a whole increased in body length following the extinction of the

carcharodontosaurids Carcharodontosauridae (carcharodontosaurids; from the Greek καρχαροδοντόσαυρος, ''carcharodontósauros'': "shark-toothed lizards") is a group of carnivorous theropod dinosaurs. In 1931, Ernst Stromer named Carcharodontosauridae ...

in the Southern Hemisphere in the early

Late 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'', the ...

. It is hypothesized that the absence of these large apex predators allowed other theropods to diversify and fill the empty niche space. Megaraptorids, in addition to abelisaurids and unenlagiids, then became the primary predators in their ecosystems. An analysis of known megaraptorans shows that members of the clade in

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

,

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

, and

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

ranged from long during the

Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...

-

Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...

, after which

Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ago), a ...

-early

Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...

forms increased to in Australia and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

.

Turonian The Turonian is, in the ICS' geologic timescale, the second age in the Late Cretaceous Epoch, or a stage in the Upper Cretaceous Series. It spans the time between 93.9 ± 0.8 Ma and 89.8 ± 1 Ma (million years ago). The Turonian is preceded by t ...

-

Coniacian The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by t ...

megaraptorids, only known from South America, were larger, at long, and the final

Santonian The Santonian is an age in the geologic timescale or a chronostratigraphic stage. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series. It spans the time between 86.3 ± 0.7 mya (million years ago) and 83.6 ± 0.7 mya. The ...

-

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...

forms were larger still, at . Traces on the preserved ribs were interpreted by the describing authors as being attachment points for ligaments. From these, it was inferred that ''Maip'' would have likely had a

respiratory system The respiratory system (also respiratory apparatus, ventilatory system) is a biological system consisting of specific organs and structures used for gas exchange in animals and plants. The anatomy and physiology that make this happen varies grea ...

comparable to those of modern

birds 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 lightweigh ...

, rather than those of other extant reptiles, such as crocodilians.

Classification

In their phylogenetic analyses, Rolando ''et al.'' (2022) recovered ''Maip'' as a derived megaraptorid in a polytomy with other Argentinian megaraptorids. They also noted the presence of two distinct clades: a more inclusive clade, comprising all megaraptorids except '' Fukuiraptor'' and '' Australovenator'', (shown below as "Clade A"), and a more exclusive clade of larger, entirely South American megaraptorids (shown below as "Clade B"). Like some previous analyses by other authors, Megaraptora is nested within Coelurosauria, as the sister taxon to

Tyrannosauroidea Tyrannosauroidea (meaning 'tyrant lizard forms') is a superfamily (or clade) of coelurosaurian theropod dinosaurs that includes the family Tyrannosauridae as well as more basal relatives. Tyrannosauroids lived on the Laurasian supercontinent b ...

. The cladogram below displays the megaraptoran results of the phylogenetic analyses by Rolando ''et al'' (2022).

Paleoecology

''Maip'' is known from the

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...

-dated

Chorrillo Formation The Chorrillo Formation, also named as Chorillo Formation,Río Leona
at Isasicursor ''Isasicursor'' (meaning "Isasi's runner" after Marcelo Pablo Isasi) is a genus of elasmarian ornithopod from the Chorrillo Formation from Santa Cruz Province in Argentina. The type and only species is ''Isasicursor santacrucensis''. It was a c ...

'', an elasmarian

ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous world ...

, and ''

Nullotitan ''Nullotitan'' (meaning "Nullo's giant", in honor of paleontologist Francisco Nullo) is a genus of lithostrotian titanosaur from the Chorrillo Formation from Santa Cruz Province in Argentina. The type and only species is ''Nullotitan glaciaris ...

'', a

titanosaurian Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thr ...

sauropod Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their bo ...

. Indeterminate remains belonging to

ankylosaurs Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...

, euiguanodontians,

hadrosaurids 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 includ ...

, noasaurids, and unenlagiids have also been recovered from the formation. Fossils of indeterminate anurans,

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 li ...

, mammals,

mosasaurs Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Greek ' meaning 'lizard') comprise a group of extinct, large marine reptiles from the Late Cretaceous. Their first fossil remains were discovered in a limestone quarry at Maastricht on the ...

,

snakes Snakes are elongated, limbless, carnivorous reptiles of the suborder Serpentes . Like all other squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales. Many species of snakes have skulls with several more joi ...

,

turtles Turtles are an order (biology), order of reptiles known as Testudines, characterized by a special turtle shell, shell developed mainly from their ribs. Modern turtles are divided into two major groups, the Pleurodira (side necked turtles) an ...

, and gastropods are also known. Very fragmentary fossil material, including teeth and a dorsal centrum, has been classified as having megaraptorid affinities, but it is too fragmentary to be assigned to ''Maip'' or any other taxon.

References

{{Taxonbar, from=Q111731074 Megaraptorans Prehistoric coelurosaurs Maastrichtian life Late Cretaceous dinosaurs of South America Cretaceous Argentina Fossils of Argentina Fossil taxa described in 2022