Aletopelta
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''Aletopelta'' (; meaning 'wanderer 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 ...
of basal
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 ...
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 Southern
California California is a state in the Western United States, located along the Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the most populous U.S. state and the 3rd largest by area. It is also the m ...
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'', ...
(upper
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, 75.5 Ma) in what is now the Point Loma Formation. The type and only species, ''Aletopelta coombsi'', is known from a partial skeleton preserving osteoderms. It was originally described in 1996 by W. P. Coombs, Jr. and T.A. Deméré before being named in 2001 by Tracy Ford and James Kirkland. ''Aletopelta'' has an estimated size of 5 metres (16 feet) and weight of 2 tonnes (4,409 lbs). 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 sever ...
formed a miniature
reef A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes— deposition of sand, wave erosion planing down rock o ...
and was scavenged upon by
invertebrates Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordat ...
and
sharks Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimorp ...
.


Discovery and naming

In 1987, construction work was done on the College Boulevard near Carlsbad at the Californian coast. While paleontologically surveying the work, Bradford Riney noted that a skeleton had been uncovered by a ditch dug for a sewage pipe. Within days, the specimen was secured by the
San Diego Natural History Museum The San Diego Natural History Museum is a museum located in Balboa Park in San Diego, California. It was founded in 1874 as the San Diego Society of Natural History. It is the second oldest scientific institution west of the Mississippi and th ...
. It was dubbed the "Carlsbad Ankylosaur".W.P. Coombs, Jr. and T.A. Deméré, 1996, "A Late Cretaceous nodosaurid ankylosaur (Dinosauria: Ornithischia) from marine sediments of coastal California", ''Journal of Paleontology'' 70(2): 311-326 The skeleton was later described, but not named, in 1996 by Thomas Deméré and Walter Preston Coombs before being named in 2001 by Tracy Lee Ford and James Kirkland.T.L. Ford and J.I. Kirkland, 2001, "Carlsbad ankylosaur (Ornithischia, Ankylosauria): an ankylosaurid and not a nodosaurid", In: ''The Armored Dinosaurs''. Indiana University Press, Bloomington pp 239-260 The skeleton originated from a layer of the marine Point Loma Formation which dates to the upper
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, 75.5 Ma. The formation has yielded specimens pertaining to calcareous nannoplankton, foraminifers, scaphopods, pelecypods, gastropods, cephalopods, ostracods, decapods, echinoids, elasmobranchs, and actinopterygians, with the addition of a
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
, right
dentary In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
containing
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, t ...
and cervical
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 ...
of a
hadrosaur 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 incl ...
. The type and only known specimen of ''Aletopelta'' was once a bloated carcass that had been washed out to sea, likely by a stream, which sank to the bottom with its underside facing upwards and became a miniature
reef A reef is a ridge or shoal of rock, coral or similar relatively stable material, lying beneath the surface of a natural body of water. Many reefs result from natural, abiotic processes— deposition of sand, wave erosion planing down rock o ...
, as evidence by Pelecypoda such as ''
Ostrea ''Ostrea'' is a genus of edible oysters, marine bivalve mollusks in the family Ostreidae, the oysters. Fossil records This genus is very ancient. It is known in the fossil records from the Permian to the Quaternary (age range: from 259 to 0. ...
sp.'' and ''
Spondylus ''Spondylus'' is a genus of bivalve molluscs, the only genus in the family Spondylidae.MolluscaBase (2019). MolluscaBase. Spondylus Linnaeus, 1758. Accessed through: World Register of Marine Species at: http://www.marinespecies.org/aphia.php?p=t ...
sp.'' being attached to the bones and the presence of
ammonites Ammonoids are a group of extinct marine mollusc animals in the subclass Ammonoidea of the class Cephalopoda. These molluscs, commonly referred to as ammonites, are more closely related to living coleoids (i.e., octopuses, squid and cuttl ...
and gastropods found in association with the skeleton. The carcass was also scavenged upon by marine invertebrates and sharks such as ''
Squalicorax ''Squalicorax'', commonly known as the crow shark, is a genus of extinct lamniform shark known to have lived during the Cretaceous period. The genus had a global distribution in the Late Cretaceous epoch. Multiple species within this genus are c ...
'' and ''
Scapanorhynchus ''Scapanorhynchus'' (from el, σκάφιου , 'shovel' and el, ῥύγχος 'snout') is an extinct genus of shark that lived from the early Cretaceous until possibly the Miocene if ''S. subulatus'' is a mitsukurinid and not a sand shark.Capet ...
'', which resulted with most of the long bones being hollow and many shallow pits on the osteoderms and ribs. 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 sever ...
specimen, SDNHM 33909, consists of
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, t ...
, 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 eith ...
e, partial humerus, partial
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 ...
, possible fragment of right
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also 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 ...
, partial left and possibly right ischium,
femora The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with t ...
,
tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...
e,
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity i ...
e, four or five partial
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 ...
, dorsal neural arch, neural arches of the sacrum, fragmentary ribs,
osteoderms Osteoderms are bony deposits forming scales, plates, or other structures based in the dermis. Osteoderms are found in many groups of extant and extinct reptiles and amphibians, including lizards, crocodilians, frogs, temnospondyls (extinct amphi ...
including
pelvic The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The p ...
shield and cervical half ring. The type specimen may represent an immature individual based on the unfused
astragalus ''Astragalus'' is a large genus of over 3,000 species of herbs and small shrubs, belonging to the legume family Fabaceae and the subfamily Faboideae. It is the largest genus of plants in terms of described species. The genus is native to tempe ...
, partly fused scutes and unfused neural spines. The generic name, ''Aletopelta'', is composed of the
Greek Greek may refer to: Greece 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 ...
words "''aletes''" (wanderer) and "''pelte''" (small shield), in reference to its armour and the
tectonic plate Plate tectonics (from the la, label=Late Latin, tectonicus, from the grc, τεκτονικός, lit=pertaining to building) is the generally accepted scientific theory that considers the Earth's lithosphere to comprise a number of large te ...
containing the Peninsular Ranges Terrane was somewhere opposite the middle of
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
, which also carried the holotype specimen with it. The specific name, coombsi, honours the vertebrate palaeontologist Walter Preston Coombs, Jr., for their work and years of research.


Description


Size and distinguishing traits

Ford & Kirkland (2001) originally gave ''Aletopelta'' an estimated length of 6 metres (19.7 feet). However,
Gregory S. Paul Gregory Scott Paul (born December 24, 1954) is an American freelance researcher, author and illustrator who works in paleontology, and more recently has examined sociology and theology. He is best known for his work and research on theropod dino ...
gave a lower estimate of 5 metres (16 feet) long and weight of 2 tonnes (4,409 lbs) in 2010. Ford and Kirkland (2001) originally diagnosed ''Aletopelta'' based on the teeth being wider than tall; the femur being much longer than both the tibia and fibula; the presence of three metatarsals; a pelvic shield composed of polygonal, low-peaked osteoderms; a large short-pointed spike in the shoulder region; the presence of hollow cap-like osteoderms across dorsum; the presence of hollow pup-tent-like osteoderms over neck and shoulders; triangular, dorsally compressed caudal osteoderms that are highly asymmetrical from top to bottom; and hollow and thin osteoderms. However, Arbour & Currie (2015) later diagnosed Aletopelta based on the presence of hexagonal pelvic osteoderms that form a semi-continuous sheet over the pelvis; the pelvic osteoderms lack a dendritic surface texture; the first cervical half ring is composed of osteoderms fused to an underlying bony band rather than being closely appressed adjacent osteoderms.


Classification

Deméré & Coombs (1996) originally interpreted SDNHM 33909 as an indeterminate
nodosaurid 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, we ...
that showed similarities with ''
Edmontonia ''Edmontonia'' is a genus of panoplosaurin nodosaurid dinosaur from the Late Cretaceous Period. It is part of the Nodosauridae, a family within Ankylosauria. It is named after the Edmonton Formation (now the Horseshoe Canyon Formation in Ca ...
'', ''
Panoplosaurus ''Panoplosaurus'' is a genus of armoured dinosaur from the Late Cretaceous of Alberta, Canada. Few specimens of the genus are known, all from the middle Campanian of the Dinosaur Park Formation, roughly 76 to 75 million years ago. It was first d ...
'' and ''
Stegopelta ''Stegopelta'' (meaning "roofed shield") is a genus of struthiosaurin nodosaurid dinosaur. It is based on a partial skeleton from the latest Albian- earliest Cenomanian-age Lower and Upper Cretaceous Belle Fourche Member of the Frontier Forma ...
'' but refrained from naming it as they considered it to be undiagnostic. Ford (2000) considered that SDNHM 33909 formed the clade Stegopeltinae, along with ''Stegopelta'' and '' Glyptodontopelta'', within
Ankylosauridae 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–Pa ...
based on the presence of a pelvic shield composed of co-ossified hexagonal osteoderms.T.L. Ford. (2000). "A review of ankylosaur osteoderms from New Mexico and a preliminary review of ankylosaur armor", In: S. G. Lucas and A. B. Heckert (eds.), ''Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science Bulletin'' 17: 157-176 Ford & Kirkland (2001) also reinterpreted it as an ankylosaurid and considered it to be taxonomically distinct enough to be warranted as a new genus and species. Vickaryous et al. (2004) considered ''Aletopelta'' as a '' nomen dubium'' due to the lack of diagnostic characteristics. Arbour & Currie (2015), however, considered it as a valid genus based on new characteristics and found Stegopeltinae to not be a valid clade. Arbour & Currie (2015) additionally recovered ''Aletopelta'' within 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 ...
containing ''
Liaoningosaurus ''Liaoningosaurus'' is an unusual genus of ankylosaurian dinosaurs from the Early Cretaceous period of China. It contains a single species, ''Liaoningosaurus paradoxus'', and is represented by two fossil specimens collected from the Yixian Form ...
'', ''
Cedarpelta ''Cedarpelta'' is a extinct genus of basal 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 we ...
'', ''
Chuanqilong ''Chuanqilong'' (meaning "legendary dragon") is a monospecific genus of basal ankylosaurid dinosaur from the Liaoning Province, China that lived during the Early Cretaceous (late Barremian to Aptian stage, 122.0 to 118.9 Ma) in what is now the ...
'' and more derived ankylosaurids. A similar position was recovered by Arbour & Evans (2017) and Rivera-Sylva et al. (2018) but also placed ''
Hylaeosaurus ''Hylaeosaurus'' ( ; Greek: / "belonging to the forest" and / "lizard") is a herbivorous ankylosaurian dinosaur that lived about 136 million years ago, in the late Valanginian stage of the early Cretaceous period of England. It was found ...
'', ''
Crichtonpelta ''Crichtonpelta'' is a genus of extinct herbivorous ankylosaurid dinosaur from the Late Cretaceous ( Cenomanian) of China. Discovery and naming In 2007, Lü Junchang, Ji Qiang, Gao Yubo and Li Zhixin named and described a second species of ''C ...
'', ''
Gobisaurus ''Gobisaurus'' is an extinct genus of herbivorous basal ankylosaurid ankylosaur from the Upper Cretaceous (and possibly also the Lower Cretaceous) of China (''Nei Mongol Zizhiqu''). The genus is monotypic, containing only the species ''Gobisa ...
'' and ''
Shamosaurus ''Shamosaurus'' is an extinct genus of herbivorous basal ankylosaurid ankylosaur from Early Cretaceous ( Aptian to Albian stage) deposits of Höövör, Mongolia. Discovery and naming In 1977, a Soviet-Mongolian expedition discovered the ske ...
'' within the polytomy. Park et al. (2019) also placed ''Aletopelta'' within a polytomy but with the inclusion of '' Jinyunpelta'', ''
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'' grew ...
'' and ''
Kunbarrasaurus ''Kunbarrasaurus'' is an extinct genus of small herbivorous ankylosaurian dinosaur from the Cretaceous of Australia. Discovery In November 1989, at Marathon Station near Richmond, Queensland, the skeleton was discovered of an ankylosauria ...
''. Although most often being recovered as a basal ankylosaurid, Frauenfelder et al. (2022) found it to be the basalmost nodosaurid. The results of a phylogenetic analysis conducted by Arbour & Currie (2015) is reproduced below.


See also

*
Timeline of ankylosaur research This timeline of ankylosaur research is a chronological listing of events in the history of paleontology focused on the ankylosaurs, quadrupedal herbivorous dinosaurs who were protected by a covering bony plates and spikes and sometimes by a club ...


References


External links

{{Taxonbar, from=Q134299 Ankylosaurids Late Cretaceous dinosaurs of North America Fossil taxa described in 2001 Taxa named by James I. Kirkland Paleontology in California Campanian genus first appearances Campanian genus extinctions Ornithischian genera