''Ahshislepelta'' (meaning "Ah-shi-sle-pah Wash shield") is a

monospecific In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...

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 ...

ankylosaur 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. ...

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

New Mexico ) , population_demonym = New Mexican ( es, Neomexicano, Neomejicano, Nuevo Mexicano) , seat = Santa Fe, New Mexico, Santa Fe , LargestCity = Albuquerque, New Mexico, Albuquerque , LargestMetro = Albuquerque metropolitan area, Tiguex , Offi ...

that lived during the

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'', ...

(late

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 ...

stage, 74.5 Ma) in what is now the Hunter Wash Member of the

Kirtland Formation The Kirtland Formation (originally the Kirtland Shale) is a sedimentary geological formation. Description The Kirtland Formation is the product of alluvial muds and overbank sand deposits from the many channels draining the coastal plain th ...

. The type and only species, ''Ahshislepelta minor'', is known only from an incomplete

postcranial skeleton Postcrania (postcranium, adjective: postcranial) in zoology and vertebrate paleontology is all or part of the skeleton apart from the skull. Frequently, fossil remains, e.g. of dinosaurs or other extinct tetrapods, consist of partial or isolated sk ...

of a small subadult or adult individual. It was named in

2011 File:2011 Events Collage.png, From top left, clockwise: a protester partaking in Occupy Wall Street heralds the beginning of the Occupy movement; protests against Libyan dictator Muammar Gaddafi, who was killed that October; a young man celebrates ...

by Michael Burns and Robert M. Sullivan. Based on the size of the humerus, ''Ahshislepelta'' is larger than ''

Pinacosaurus mephistocephalus ''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 ...

'' but smaller than ''

Talarurus ''Talarurus'' ( ; meaning "basket tail" or "wicker tail") is a genus of ankylosaurid dinosaur that lived in Asia during the Late Cretaceous period, about 96 million to 89 million years ago. The first remains of ''Talarurus'' were discovered in ...

'' and ''

Pinacosaurus grangeri ''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 ...

''.

Discovery and naming

In 2005, a partial postcranial

skeleton A skeleton is the structural frame that supports the body of an animal. There are several types of skeletons, including the exoskeleton, which is the stable outer shell of an organism, the endoskeleton, which forms the support structure inside ...

of an

was discovered from the Hunter Wash Member in the lower part of the Kirtland Formation, New Mexico at the Ah-shi-sle-pah Wilderness Study Area. The specimen was later collected over consecutive field seasons from 2005 to 2009 and was subsequently described in 2011 by Michael Burns and Robert M. Sullivan. The

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 seve ...

specimen, SMP VP-1930, consists of a partial

girdle A belt, especially if a cord or rope, is called a girdle if it is worn as part of Christian liturgical vestments, or in certain historical, literary or sports contexts. Girdles are used to close a cassock in Christian denominations, including th ...

, partial

scapulocoracoid The scapulocoracoid is the unit of the pectoral girdle that contains the coracoid and scapula. The coracoid itself is a beak-shaped bone that is commonly found in most vertebrates with a few exceptions. The scapula is commonly known as the ''shoulde ...

, a proximal portion of the

radius In classical geometry, a radius (plural, : radii) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', ...

, numerous cervical and/or

dorsal vertebrae In vertebrates, thoracic vertebrae compose the middle segment of the vertebral column, between the cervical vertebrae and the lumbar vertebrae. In humans, there are twelve thoracic vertebrae and they are intermediate in size between the cervical ...

fragments, complete and fragmentary thoracic osteoderms, and other unidentifiable postcranial fragments. The holotype specimen is currently housed at the

State Museum of Pennsylvania The State Museum of Pennsylvania is a non-profit museum at 300 North Street in downtown Harrisburg, Pennsylvania. It is run by the state through the Pennsylvania Historical and Museum Commission and was created to preserve and interpret the region ...

Harrisburg Harrisburg is the capital city of the Commonwealth of Pennsylvania, United States, and the county seat of Dauphin County. With a population of 50,135 as of the 2021 census, Harrisburg is the 9th largest city and 15th largest municipality in ...

. The generic name, ''Ahshislepelta'', is derived from Ah-shi-sle-pah Wash, the locality from which the holotype came, and the

Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...

word "''peltè''" (shield). The specific name, minor, is in reference to its small adult size relative to other

North American North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the ...

ankylosaurids. The holotype specimen represents a subadult or adult individual based on the complete fusion of the scapulocoracoids, centra and

neural arches 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 characteristic i ...

of the vertebrae, and the re-modelled osteoderm cores. The holotype This new specimen falls within the same size range as adult specimens of other

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 ...

taxa outside of North America and represents an individual that was larger than juvenile specimens of ''Pinacosaurus grangeri'' and ''Pinacosaurus mephistocephalus'' but smaller than Talarurus and adult specimens of ''P. grangeri'' based on humeral size; however, the scapulocoracoid is larger than that of ''Talarurus''.

Description

Distinguishing traits

Burns & Sullivan (2011) diagnosed ''Ahshislepelta'' based on the dorsolateral overhang of the scapular acromion process to 25% of the dorsoventral width of the

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 eith ...

. ''Ahshislepelta'' differs from other ankylosaurids, with the exception of ''

Euoplocephalus ''Euoplocephalus'' ( ) is a genus of very large, herbivorous ankylosaurid dinosaurs, living during the Late Cretaceous of Canada. It has only one named species, ''Euoplocephalus tutus''. The first fossil of ''Euoplocephalus'' was found in 1 ...

'', by the superficial osteodermal surface texture, characterized by uniformly distributed pitted rugosity, and sparse distribution of reticular neurovascular grooves with neurovascular foramina extending perpendicularly to obliquely into the bone.

Postcrania

The right scapula preserves osteoderms on the lateral side at the most distal portion and is associated with fragments of

ribs The rib cage, as an enclosure that comprises the ribs, vertebral column and sternum in the thorax of most vertebrates, protects vital organs such as the heart, lungs and great vessels. The sternum, together known as the thoracic cage, is a semi- ...

that are mostly visible on the medial side. The scapula is fully fuse to the

coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) 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 prese ...

and has a blunt, rugose process along the anterodorsal margin for the attachment of

muscles Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of musc ...

. The coracoid is roughly 16 cm long and the coracoid foramen is anterodorsal to the glenoid cavity. The glenoid cavity forms an arc that projects the scapular margin posteroventrally at an 80° angle relative to the coracoid margin. As in

nodosaurids Nodosauridae is a family of ankylosaurian dinosaurs, from the Late Jurassic to the Late Cretaceous period in what is now North America, South America, Europe, and Asia. Description Nodosaurids, like their close relatives the ankylosaurids, were ...

, the scapula does not narrow towards the upper surface and sides relative to the coracoid and does not exhibit a scapular neck. A rugose, ridge-like structure that is identified as the

acromion In human anatomy, the acromion (from Greek: ''akros'', "highest", ''ōmos'', "shoulder", plural: acromia) is a bony process on the scapula (shoulder blade). Together with the coracoid process it extends laterally over the shoulder joint. The ...

is present along the dorsal margin of the scapula opposite the glenoid cavity. The acromion attains its front most extension towards the back and sides to the most anterior extent of the glenoid cavity. Due to

taphonomic Taphonomy is the study of how organisms decay and become fossilized or preserved in the paleontological record. The term ''taphonomy'' (from Greek , 'burial' and , 'law') was introduced to paleontology in 1940 by Soviet scientist Ivan Efremov t ...

shearing, the prominent overhang of the acromion on the left scapula is absent, although this feature is not a taphonomic artefact on the right scapulocoracoid. However, the right scapulocoracoid is cracked, its junction with the scapular blade is visible, well-preserved, and verifies its natural projection towards the lower edge and sides. Towards the sides of the acromion, the borders of the

infraspinous fossa The infraspinous fossa (infraspinatus fossa or infraspinatous fossa) of the scapula is much larger than the supraspinatous fossa; toward its vertebral margin a shallow concavity is seen at its upper part; its center presents a prominent convexi ...

are prominent and creates an area or the attachment of muscles. The left forelimb is more than 50% complete. As in ankylosaurids, the left

humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a r ...

is massive and has a length of 31 cm. The left humerus preserves a deltopectoral crest that is well-developed and measures 15 cm across the widest portion of the humerus. The lateral margin of the deltopectoral crest forms a 23° angle in posterior view with the long axis of the humerus. Bounded by the deltopectoral crest and humeral head is a broad, bicipital fossa that is present on the anterior face of the humerus. On the anterior face of the humerus is a hemispherical radial condyle. The proximal articular surface of the left

is oval in proximal view and is concave. The articular surface is 8 cm long, proportionally twice the diameter of the

diaphysis The diaphysis is the main or midsection (shaft) of a long bone. It is made up of cortical bone and usually contains bone marrow and adipose tissue (fat). It is a middle tubular part composed of compact bone which surrounds a central marrow cavit ...

. Numerous

vertebrae 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 ...

fragments that probably represent parts of the cervicals and/or dorsals were recovered and the position of the vertebrae are based on their association with elements of the pectoral girdles and forelimb. A few vertebrae were dorsoventrally tall dorsal vertebrae, based on a few of the more complete vertebrae appearing to have laterally compressed neural canals. All other vertebrae lack open sutural facets for the neural arches.

Osteoderms

Numerous osteoderms and associated osteoderm fragments were associated with the holotype specimen. Most osteoderms pertain to the thoracic region while others might pertain to the pelvic region and

forelimbs A forelimb or front limb is one of the paired articulated appendages (limbs) attached on the cranial (anterior) end of a terrestrial tetrapod vertebrate's torso. With reference to quadrupeds, the term foreleg or front leg is often used instead. ...

. All osteoderms are either keeled or circular with an off-center apex. The distal portion of the right scapula preserves an ''in situ'' osteoderm and numerous ossicles. The osteoderm and ossicles of the right scapula and another series of osteoderms preserved in situ display a rosette arrangement of small ossicles surrounding the osteoderms. The largest osteoderm pertains to the thoracic region behind the cervical region based on the similar morphology to the median thoracic osteoderms of the holotype specimen of Scolosaurus. The osteoderm has a length of 15 cm, a width of 12 cm and a height of 10 cm. The smallest ossicle has a diameter of 1 cm. The surface texture of the osteoderms are smooth to uniformly pitted with a sparse patterns of grooves as in ''Euoplocephalus''. One osteoderm has a histological condition typical of that of ankylosaurids.

Classification

Burns & Sullivan (2011) originally placed ''Ahshislepelta'' as an ankylosaurid within the clade

Ankylosaurinae Ankylosaurinae is a subfamily of ankylosaurid dinosaurs, existing from the Early Cretaceous about 105 million years ago until the end of the Late Cretaceous, about 66 mya. Many genera are included in the clade, such as ''Ankylosaurus'', ''Pina ...

, although this was not based on the result of 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 ...

. Likewise, Arbour & Currie (2015) placed ''Ahshislepelta'' as sister taxon to '' Gastonia'' at the base of Ankylosauridae, and Wiersma and Irmis (2018) placed it in a polytomy consisting of ''

'', '' Oohkotokia'', ''

Scolosaurus ''Scolosaurus'' is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the lower levels of the Dinosaur Park Formation and upper levels of the Oldman Formation in the Late Cretaceous (latest middle Cam ...

'', ''

Ziapelta ''Ziapelta'' is an extinct genus of ankylosaurid. Its fossils have been found in the Hunter Wash and De-na-zin members of the Kirtland Formation of Upper Cretaceous (Campanian) New Mexico. It was named in 2014, in a research paper led by ankylos ...

'', ''

Anodontosaurus ''Anodontosaurus'' is an extinct genus of ankylosaurid dinosaurs within the subfamily Ankylosaurinae. It is known from the entire span of the Late Cretaceous Horseshoe Canyon Formation (mid Late Campanian to "middle" Maastrichtian stage, about 7 ...

'', ''

Ankylosaurus ''Ankylosaurus'' is a genus of armored dinosaur. Its fossils have been found in geological formations dating to the very end of the Cretaceous Period, about 68–66 million years ago, in western North America, making it among the last of th ...

'' and '' Dyoplosaurus''. On the other hand, several other studies have favored a nodosaurid position. Arbour ''et al.'' (2016) recovered it 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 tr ...

with ''

Niobrarasaurus ''Niobrarasaurus'' (meaning "Niobrara lizard") is an extinct genus of nodosaurid ankylosaur which lived during the Cretaceous 87 to 82 million years ago. Its fossils were found in the Smoky Hill Chalk Member of the Niobrara Formation, in western ...

'' and a juvenile nodosaurid from the

Paw Paw Formation The Paw Paw Formation is a geological formation in Texas whose strata date back to the late Albian stage of the Early Cretaceous. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, David B; et al. (20 ...

, while Zheng ''et al.'' (2018) positioned ''Ahshislepelta'' in a polytomy containing '' Tatankacephalus'', ''

Silvisaurus ''Silvisaurus'', from the Latin silva "woodland" and Greek sauros "lizard", is a nodosaurid ankylosaur from the Early to Late Cretaceous period. Discovery and species A fossil of the species was discovered in the fifties by rancher Warren ...

'', ''Niobrarasurus'', ''

Nodosaurus ''Nodosaurus'' (meaning "knobbed lizard") is a genus of herbivorous nodosaurid ankylosaurian dinosaur from the Late Cretaceous, the fossils of which are found exclusively in the Frontier Formation in Wyoming. Description ''Nodosaurus'' gre ...

'' and more deeply nested taxa. Rivera-Sylva ''et al.'' (2018) placed it as sister taxon to ''Niobrarasurus''. A phylogenetic analysis conducted by Rivera-Sylva ''et al.'' (2018) is reproduced below, favoringa nodosaurid position. Clade names have been added following Madzia ''et al.'' (2021). In contrast, an earlier analysis by Arbour & Currie (2015) favors an ankylosaurid position. Their results are reproduced below.

Paleoenvironment

''Ahshislepelta'' is known from the Hunter Wash Member of the Kirtland Formation which has been dated to the upper

stage, 74.5 Ma. The Kirtland Formation consists of interbedded

sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicat ...

siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, ...

mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology.' ...

coal Coal is a combustible black or brownish-black sedimentary rock, formed as rock strata called coal seams. Coal is mostly carbon with variable amounts of other elements, chiefly hydrogen, sulfur, oxygen, and nitrogen. Coal is formed when ...

and

shale Shale is a fine-grained, clastic sedimentary rock formed from mud that is a mix of flakes of clay minerals (hydrous aluminium phyllosilicates, e.g. kaolin, Al2 Si2 O5( OH)4) and tiny fragments (silt-sized particles) of other minerals, especiall ...

. The Kirtland Formation lays on the western margin of the

Western Interior Seaway The Western Interior Seaway (also called the Cretaceous Seaway, the Niobraran Sea, the North American Inland Sea, and the Western Interior Sea) was a large inland sea that split the continent of North America into two landmasses. The ancient sea ...

and represents a

floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river which stretches from the banks of its channel to the base of the enclosing valley walls, and which experiences flooding during periods of high discharge.Goudi ...

that was abundant with

ferns A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except th ...

conifers Conifers are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All ext ...

and

flowering plants Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...

. Based on the abundance of angiosperms with leaves that have entire or nearly entire margins and drip points, the Kirtland Formation may have had a warm temperate to subtropical climate. The Kirtland Formation was better drained than the underlying

Fruitland Formation The Fruitland Formation is a geologic formation found in the San Juan Basin in the states of New Mexico and Colorado, in the United States of America. It contains fossils dating it to the Campanian age of the late Cretaceous.

due to the lack of coal swamps. ''Ahshislepelta'' coexisted with the

pachycephalosaurid Pachycephalosauria (; from Greek παχυκεφαλόσαυρος for 'thick headed lizards') is a clade of ornithischian dinosaurs. Along with Ceratopsia, it makes up the clade Marginocephalia. With the exception of two species, most pachycephal ...

Stegoceras ''Stegoceras'' is a genus of pachycephalosaurid (dome-headed) dinosaur that lived in what is now North America during the Late Cretaceous period, about 77.5 to 74 million years ago (mya). The first specimens from Alberta, Canada, were descri ...

'', the chasmosaurine

ceratopsids Ceratopsidae (sometimes spelled Ceratopidae) is a family of ceratopsian dinosaurs including ''Triceratops'', ''Centrosaurus'', and ''Styracosaurus''. All known species were quadrupedal herbivores from the Upper Cretaceous. All but one species ar ...

Navajoceratops ''Navajoceratops'' (meaning "Navajo horned face") is a genus of ceratopsid dinosaur from the late Cretaceous Period of what is now North America. The genus contains a single species, ''N. sullivani'', named after Robert M. Sullivan, leader of th ...

'' and ''

Terminocavus ''Terminocavus'' is a genus of ceratopsid dinosaur from the late Cretaceous Period of what is now North America. The genus contains a single species, the type species ''Terminocavus sealeyi'', known from a parietal and some other associated frag ...

'', the ankylosaurine ankylosaurid ''

'', the eutyrannosaur

tyrannosauroid 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 ...

Bistahieversor ''Bistahieversor'' (meaning "Bistahi destroyer"), also known as the "Bisti Beast", is a genus of eutyrannosaurian tyrannosauroid dinosaur; the genus contains only a single known species, ''B. sealeyi'', described in 2010, from the Late Cretaceo ...

'', and the

azhdarchid Azhdarchidae (from the Persian word , , a dragon-like creature in Persian mythology) is a family of pterosaurs known primarily from the Late Cretaceous Period, though an isolated vertebra apparently from an azhdarchid is known from the Early Cre ...

pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...

Navajodactylus ''Navajodactylus'' (meaning "Navajo finger") is an extinct genus of pterodactyloid pterosaur from Late Cretaceous (late Campanian stage) deposits of the San Juan Basin in New Mexico, United States. Discovery The holotype specimen of ''Navajodac ...

''.

References

{{Taxonbar, from=Q1705974 Late Cretaceous dinosaurs of North America Ankylosaurs Fossil taxa described in 2011 Paleontology in New Mexico Campanian genus first appearances Campanian genus extinctions Ornithischian genera

Discovery and naming

Description

Distinguishing traits

Postcrania

Osteoderms

Classification

Paleoenvironment

See also

References