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''Maip'' is a genus of large megaraptorid
theropod Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur from the
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
(
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
)
Chorrillo Formation The Chorrillo Formation, also named as Chorillo Formation,Río Leona
at Fossilworks.org is ...
of Santa Cruz,
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. 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 considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
, and possibly the world.


Discovery and naming

The ''Maip''
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
specimen, MPM 21545, was discovered by Alexis Rolando on the La Anita Farm, west of
El Calafate El Calafate, also called ''Calafate'', is a city in the Argentine province of Santa Cruz, in Patagonia. It is located on the southern border of Lago Argentino, in the southwest part of the province (about northwest of Río Gallegos). The name ...
, 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 An axis (: axes) may refer to: Mathematics *A specific line (often a directed line) that plays an important role in some contexts. In particular: ** Coordinate axis of a coordinate system *** ''x''-axis, ''y''-axis, ''z''-axis, common names ...
, several
dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...
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 th ...
vertebrae, cervical and dorsal
ribs The rib cage or thoracic cage is an endoskeletal enclosure in the thorax of most vertebrates that comprises the ribs, vertebral column and sternum, which protect the vital organs of the thoracic cavity, such as the heart, lungs and great vessels ...
,
gastralia Gastralia (: gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In these reptil ...
, a left
coracoid A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...
, fragmentary
scapula The scapula (: 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 either side ...
, partial right pubis, and partial
metatarsal The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
. 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 versi ...
in 2021. However, since the paper did not meet the necessary
requirements In engineering, a requirement is a condition that must be satisfied for the output of a work effort to be acceptable. It is an explicit, objective, clear and often quantitative description of a condition to be satisfied by a material, design, pro ...
, 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, the Tehuelche people, were nomadic hunters who occupied territory in present-day Chile, north of Tierra del Fuego and south of th ...
mythology that is "the shadow of death" that "kills with cold wind." The specific name, "''macrothorax''", is derived from the
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
"makrós", meaning "large", and the
Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
"thorax", meaning "chest", in reference to its large
thoracic cavity The thoracic cavity (or chest cavity) is the chamber of the body of vertebrates that is protected by the thoracic wall (rib cage and associated skin, muscle, and fascia). The central compartment of the thoracic cavity is the mediastinum. Ther ...
.


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 in the Southern Hemisphere in the early
Late Cretaceous The Late Cretaceous (100.5–66 Ma) is the more recent 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 ''cre ...
. It is hypothesized that the absence of these large
apex predator An apex predator, also known as a top predator or superpredator, is a predator at the top of a food chain, without natural predators of its own. Apex predators are usually defined in terms of trophic dynamics, meaning that they occupy the hig ...
s allowed other theropods to diversify and fill the empty niche space. Megaraptorids, in addition to
abelisaurids Abelisauridae (meaning "Abel's lizards") is a family (biology), family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous Period (geology), period, on the ancient southern supercontinent of Gondwana, and ...
and
unenlagiids 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 gro ...
, then became the primary predators in their ecosystems. An analysis of known megaraptorans shows that members of the clade in
Asia Asia ( , ) is the largest continent in the world by both land area and population. It covers an area of more than 44 million square kilometres, about 30% of Earth's total land area and 8% of Earth's total surface area. The continent, which ...
,
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
, and
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
ranged from long during the
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 125.77 Ma (million years ago) and 121.4 ± 1.0 Ma (Historically, this stage was placed at 129.4 million to approximately 125 million years ago) It is a ...
-
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
, after which
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
-early
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
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 considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
.
Turonian The Turonian is, in the International Commission on Stratigraphy, ICS' geologic timescale, the second age (geology), age in the Late Cretaceous epoch (geology), Epoch, or a stage (stratigraphy), stage in the Upper Cretaceous series (stratigraphy), ...
-
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 ...
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 m ...
-
Maastrichtian The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
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
bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...
s, rather than those of other extant reptiles, such as
crocodilia Crocodilia () is an order of semiaquatic, predatory reptiles that are known as crocodilians. They first appeared during the Late Cretaceous and are the closest living relatives of birds. Crocodilians are a type of crocodylomorph pseudosuchia ...
ns.


Classification

In their
phylogenetic analyses In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as Computational phylogenetics, phylogenetic inference. It infers the relationship among organ ...
, Rolando ''et al.'' (2022) recovered ''Maip'' as a derived megaraptorid in a
polytomy An internal node of a phylogenetic tree is described as a polytomy or multifurcation if (i) it is in a rooted tree and is linked to three or more child subtrees or (ii) it is in an unrooted tree and is attached to four or more branches. A tree ...
with other Argentinian megaraptorids. They also noted the presence of two distinct clades: a more inclusive clade, comprising all megaraptorids except ''
Fukuiraptor ''Fukuiraptor'' ("thief of Fukui") is a genus of medium-sized megaraptoran theropod dinosaur of the Early Cretaceous epoch (either Barremian or Aptian) that lived in what is now Japan. ''Fukuiraptor'' is known from the Kitadani Formation and ...
'' and ''
Australovenator ''Australovenator'' (meaning "southern hunter") is a genus of megaraptoran theropod dinosaur from Cenomanian (Late Cretaceous)-age Winton Formation (dated to 95 million years ago) of Australia. Some specimens from the Albian-aged Eumeralla Forma ...
'', (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 Megaraptora is a clade of carnivorous theropod dinosaurs. Its derived members, the Megaraptoridae are noted for their large hand claws and powerfully-built forelimbs, which are usually reduced in size in other large theropods. Although undoubt ...
is nested within
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, tyra ...
, 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 ...
. 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 International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
-dated
Chorrillo Formation The Chorrillo Formation, also named as Chorillo Formation,Río Leona
at Fossilworks.org is ...
of southern Argentina. Other named dinosaurs known from the formation include '' Isasicursor'', an
elasmaria Elasmaria is a clade of Ornithopoda, ornithopods known from Cretaceous deposits in the former Gondwana (South America, Antarctica, Australia, and possibly Africa) that contains many bipedal ornithopods that were previously considered Hypsilophodo ...
n
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
, and '' Nullotitan'', a
titanosauria Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of Sauropoda, sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with tax ...
n
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 b ...
. Indeterminate remains belonging to
ankylosaurs Ankylosauria is a group of herbivorous dinosaurs of the clade Ornithischia. It includes the great majority of dinosaurs with Armour (zoology), armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short ...
, euiguanodontians,
hadrosaurids Hadrosaurids (), also hadrosaurs 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 fami ...
, noasaurids, and
unenlagiids 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 gro ...
have also been recovered from the formation. Fossils of indeterminate
anurans A frog is any member of a diverse and largely semiaquatic group of short-bodied, tailless amphibian vertebrates composing the order Anura (coming from the Ancient Greek , literally 'without tail'). Frog species with rough skin texture due to ...
,
fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...
,
mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...
s,
mosasaur Mosasaurs (from Latin ''Mosa'' meaning the 'Meuse', and Ancient Greek, Greek ' meaning 'lizard') are an extinct group of large aquatic reptiles within the family Mosasauridae that lived during the Late Cretaceous. Their first fossil remains wer ...
s,
snake Snakes are elongated limbless reptiles of the suborder Serpentes (). Cladistically squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales much like other members of the group. Many species of snakes have s ...
s,
turtle Turtles are reptiles of the order (biology), order 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) and Crypt ...
s, and
gastropods Gastropods (), commonly known as slugs and snails, belong to a large taxonomic class of invertebrates within the phylum Mollusca called Gastropoda (). This class comprises snails and slugs from saltwater, freshwater, and from the land. Ther ...
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 Megaraptora Dinosaur genera Maastrichtian dinosaurs Fossil taxa described in 2022 Dinosaurs of Argentina