Jakapil
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''Jakapil'' (meaning "shield bearer" in Puelchean) is a genus of basal thyreophoran dinosaur from the
Candeleros Formation The Candeleros Formation is a geologic formation that outcrop, crops out in the Rio Negro Province, Río Negro, Neuquen Province, Neuquén, and Mendoza Province, Mendoza provinces of northern Patagonia, Argentina. It is the oldest formation in the ...
of Argentina. The type species is ''Jakapil kaniukura''.


Discovery and naming

The
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 ...
, MPCA-PV-630, is a partial skeleton including several
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of Extant taxon, extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, Temnospondyli, ...
and a complete lower jaw, which were found on land owned by the Mariluan family in 2012 and excavated between 2014 and 2019/2020. The finds were prepared by L. Pazo and J. Kaluza. The generic name, "''Jakapil''", is derived from "Ja-Kapïl", a Puelchean word meaning "shield bearer". This is also the literal meaning of the clade name
Thyreophora Thyreophora ("shield bearers", often known simply as "armored dinosaurs") is a group of armored ornithischian dinosaurs that lived from the Early Jurassic until the end of the Cretaceous. Thyreophorans are characterized by the presence of bod ...
. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
, "''kaniukura''", means "crest stone" in
Mapudungun Mapuche ( , ; from 'land' and 'people', meaning 'the people of the land') or Mapudungun (from 'land' and 'speak, speech', meaning 'the speech of the land'; also spelled Mapuzugun and Mapudungu) is either a language isolate or member of the s ...
, in reference to its uniquely deep jaw.


Description

''Jakapil'' represents a novel morphotype among thyreophorans, including, among other things, the presence of a
predentary Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Ancient ...
bone (absent or cartilaginous in other basal thyreophorans), large, low osteoderms, and a bipedal stance, similar to '' Scutellosaurus''. Its describers estimate it to be less than long and in weight, based on femoral circumference. While the holotype remains do not represent a juvenile individual, it was likely not fully grown and may not have been sexually mature.


Classification

Due to a combination of features seen in basal
ornithischians Ornithischia () is an extinct clade of mainly herbivorous dinosaurs characterized by a pelvic structure superficially similar to that of birds. The name ''Ornithischia'', or "bird-hipped", reflects this similarity and is derived from the Greek st ...
, basal thyreophorans, and ankylosaurs, a
phylogenetic analysis In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...
placed it, according to most data matrices, as a basal thyreophoran, outside the clade Eurypoda. Riguetti and colleagues (2022) suggested that ''Jakapil'' represents a member of a previously unknown thyreophoran clade. This clade may have diverged from the rest of the thyreophorans during the
Sinemurian In the geologic timescale, the Sinemurian is an age (geology), age and stage (stratigraphy), stage in the Early Jurassic, Early or Lower Jurassic epoch (geology), Epoch or series (stratigraphy), Series. It spans the time between 199.5 ±0.3 annu ...
. Its discovery in 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 ...
implies the existence of a long basal thyreophoran
ghost lineage A ghost lineage is a hypothesized ancestor in a species lineage that has left no fossil evidence, but can still be inferred to exist or have existed because of gaps in the fossil record or genomic evidence. The process of determining a ghost line ...
. The status of ''Jakapil'' as a thyreophoran has been disputed by some researchers; a study focused on the phylogenetic relationships of the Thyreophora noted that the only thyreophoran characteristic it possessed was dermal armor, while presenting features not present in any other thyreophoran. The premaxilla of ''Jakapil'' had no teeth, a feature not known in any thyreophoran, while the shape and depth of the dentary are more similar to those of basal neoceratopsians. Additionally, the teeth of ''Jakapil'' have a morphology distinct from any other ornithischian. Therefore, the study concluded that ''Jakapil'' most likely represents an armored marginocephalian or a member of a yet to be known lineage of non-thyreophoran ornithischians. In their 2024 analyses, Fonseca and colleagues recovered ''Jakapil'' as either the sister taxon to Eurypoda or as the basalmost member of
Ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the clade 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 ...
. They argued that ''Jakapil'' is likely not a
ceratopsia Ceratopsia or Ceratopia ( or ; Ancient Greek, Greek: "horned faces") is a group of herbivore, herbivorous, beaked dinosaurs that thrived in what are now North America, Asia and Europe, during the Cretaceous Period (geology), Period, although ance ...
n or marginocephalian, since this would required 16 additional steps. However, a position within
Parankylosauria Parankylosauria is a group of Basal (phylogenetics), basal ankylosaurian dinosaurs known from the Cretaceous of South America, Antarctica, and Australia. It is thought the group split from other ankylosaurs during the mid-Jurassic period, despite ...
only required four additional steps. As such, the authors suggested that this possibility should not be disregarded, even though it is not the most parsimonious position for the taxon.


Paleobiology

In light of its straight, narrow snout, Riguetti and colleagues (2022) suggested that ''Jakapil'' did not use its teeth and jaws to shear leaves, but instead likely processed tough plant material via
mastication Chewing or mastication is the process by which food is comminution, crushed and ground by the teeth. It is the first step in the process of digestion, allowing a greater surface area for digestive enzymes to break down the foods. During the mast ...
, as evidenced by the high amount of wear on the teeth.


Paleoenvironment

The
Candeleros Formation The Candeleros Formation is a geologic formation that outcrop, crops out in the Rio Negro Province, Río Negro, Neuquen Province, Neuquén, and Mendoza Province, Mendoza provinces of northern Patagonia, Argentina. It is the oldest formation in the ...
is interpreted as an ancient
desert A desert is a landscape where little precipitation occurs and, consequently, living conditions create unique biomes and ecosystems. The lack of vegetation exposes the unprotected surface of the ground to denudation. About one-third of the la ...
known as the Kokorkom desert, with some
oases In ecology, an oasis (; : oases ) is a fertile area of a desert or semi-desert environmentfossils A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
represent a faunal assemblage typical of Middle Cretaceous ecosystems in the western remnants of
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
. Other animals from this formation include the rhynchocephalians '' Tika'' and '' Priosphenodon'', the snake '' Najash'', the
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 ...
'' Cronopio dentiacutus'', the theropods '' Alnashetri'', ''
Buitreraptor ''Buitreraptor'' (meaning "La Buitrera seizer") is a genus of dromaeosaurid dinosaurs that lived during the Late Cretaceous of Argentina at the Candeleros Formation. ''Buitreraptor'' was described in 2005 and the type species is ''Buitreraptor g ...
'', ''
Ekrixinatosaurus ''Ekrixinatosaurus'' ('explosion-born reptile') is a genus of abelisaurid theropod which lived approximately 100 to 97 million years ago during the Late Cretaceous period. Its fossils have been found in Argentina. Only one species is currently ...
,'' and ''
Giganotosaurus ''Giganotosaurus'' ( ) is a genus of large theropod dinosaur that lived in what is now Argentina, during the early Cenomanian age of the Late Cretaceous period (geology), period, approximately 99.6 to 95 million years ago. The holotype specim ...
'', as well as the sauropods ''
Andesaurus ''Andesaurus'' ( ; "Andes lizard") is a genus of basal titanosaurian sauropod dinosaur which existed during the middle of the Cretaceous Period in South America. Like most sauropods, it would have had a small head on the end of a long neck and a ...
'' and '' Limaysaurus''.


References

{{Taxonbar, from1=Q113494797 Thyreophora Dinosaur genera Cenomanian dinosaurs Candeleros Formation Fossil taxa described in 2022 Dinosaurs of Argentina