Dracopelta
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''Dracopelta'' (meaning “
dragon A dragon is a reptilian legendary creature that appears in the folklore of many cultures worldwide. Beliefs about dragons vary considerably through regions, but dragons in western cultures since the High Middle Ages have often been depicted as ...
shield”) is a monospecific
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
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
Portugal Portugal, officially the Portuguese Republic ( pt, República Portuguesa, links=yes ), is a country whose mainland is located on the Iberian Peninsula of Southwestern Europe, and whose territory also includes the Atlantic archipelagos of ...
that lived during the
Late Jurassic The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the name ...
(uppermost lower
Tithonian In the geological timescale, the Tithonian is the latest age of the Late Jurassic Epoch and the uppermost stage of the Upper Jurassic Series. It spans the time between 152.1 ± 4 Ma and 145.0 ± 4 Ma (million years ago). It is preceded by the K ...
-upper Tithonian, 152.1-145.0 Ma) in what is now the
Lourinhã Formation The Lourinhã Formation () is a fossil rich geological formation in western Portugal, named for the municipality of Lourinhã. The formation is mostly Late Jurassic in age (Kimmeridgian/Tithonian), with the top of the formation extending into the ...
. The type and only species is ''Dracopelta zbyszewskii'', which is represented by a partial skeleton including unpublished material.


Discovery and naming

In 1963 or, more likely, early 1964, a partial
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 ...
was discovered during road construction works between the village of Barril and Praia da Assenta. Leonel Trindade confirmed the presence of the
fossil 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 ...
in the area and photographed the specimen ''in situ''. Georges Zbyszewski and Octávio da Veiga Ferreir organized the excavation and extraction of the specimen in December 1964. Parts of the specimen were unprepared and misplaced, mixed in with a specimen of ''
Miragaia Miragaia () is a former Freguesia, civil parish in the municipality of Porto, Portugal. In 2013, the parish merged into the new parish Cedofeita, Santo Ildefonso, Sé, Miragaia, São Nicolau e Vitória. The population in 2011 was 2,067, in an area ...
''. 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 several ...
specimen, MG 3 (formerly IGM 5787 and IGM 3), consists of dorsal
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 characteristic i ...
, articulated proximal
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- ...
,
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 ...
, an incomplete autopodium and unpublished material including a
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 ...
,
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 with ...
, and an ossified
tendon A tendon or sinew is a tough, high-tensile-strength band of dense fibrous connective tissue that connects muscle to bone. It is able to transmit the mechanical forces of muscle contraction to the skeletal system without sacrificing its ability ...
. Currently new material pertaining to the holotype specimen is being described. The generic name, ''Dracopelta'', is derived from the Latin word ''draco'' (dragon) and the Greek word πέλτη, pelte, (Latinised to pelta), "small shield". The specific name, ''zbyszewskii'', honours Georges Zbyszewski, after his research on fossil
vertebrates Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, ...
from Portugal. The holotype was originally, and incorrectly, identified to be from the
Kimmeridgian In the geologic timescale, the Kimmeridgian is an age in the Late Jurassic Epoch and a stage in the Upper Jurassic Series. It spans the time between 157.3 ± 1.0 Ma and 152.1 ± 0.9 Ma (million years ago). The Kimmeridgian follows the Oxfordian ...
stage of Ribamar. However, Antunes and Mateus (2003) noted that at a locality named Ribamar, there are outcrops of
Lower Cretaceous Lower may refer to: *Lower (surname) *Lower Township, New Jersey *Lower Receiver (firearms) *Lower Wick Lower Wick is a small hamlet located in the county of Gloucestershire, England. It is situated about five miles south west of Dursley, eight ...
that date to the
Valanginian In the geologic timescale, the Valanginian is an age or stage of the Early or Lower Cretaceous. It spans between 139.8 ± 3.0 Ma and 132.9 ± 2.0 Ma (million years ago). The Valanginian Stage succeeds the Berriasian Stage of the Lower Cretaceou ...
and
Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0.9 M ...
stages and deemed it as highly unlikely that the specimen was obtained from that locality. Pereda-Suberbiola et al. (2005) determined that the specimen was from a locality constrained to the uppermost lower Tithonian-upper Tithonian which shows characteristics consistent with what is recognized in the Assenta Member of the
Lourinhã Formation The Lourinhã Formation () is a fossil rich geological formation in western Portugal, named for the municipality of Lourinhã. The formation is mostly Late Jurassic in age (Kimmeridgian/Tithonian), with the top of the formation extending into the ...
, an observation agreed upon by Russo and Mateus (2021). In 2019, a specimen of an ankylosaur consisting of a nearly complete
skull The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, articulated dorsal vertebrae, proximal half of ribs, articulated anterior caudal vertebrae, mostly complete and articulated synsacrum, fragments of disarticulated and broken ribs, femora, partial with attached pelvic shield, humerus, 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 ...
, tendons mostly attached to vertebrae, and osteoderms from the Lourinhã Formation was reported. The specimen is currently being studied on to clarify if it represents an additional, more complete specimen, of ''Dracopelta'' or a new taxon.


Description

''Dracopelta'' has an estimated body length of 3 metres (9.9 feet) and a weight of 300 kilograms (600 lb).Paul, G.S., 2010, ''The Princeton Field Guide to Dinosaurs'', Princeton University Press p. 231 The holotype specimen represents an adult individual. Galton (1980) originally diagnosed ''Dracopelta'' based on the small flat osteoderms, small medial paired circular plated with raised centre and rims, long anterolateral plates, narrow nonprojecting overlapping dorsolateral plates and overlapping laterally projecting lateral plates in the thoracic region. Suberbiola et al. (2005) later diagnosed ''Dracopelta'' based on the presence of proximal
phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
II and III as long as wide and distinctive osteoderm morphology. The
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 * Dorsal co ...
ribs are gently curved which indicates that the back was broad, much like that of other ankylosaurs, ''
Dryosaurus ''Dryosaurus'' ( , meaning 'tree lizard', Greek ' () meaning 'tree, oak' and () meaning 'lizard'; the name reflects the forested habitat, not a vague oak-leaf shape of its cheek teeth as is sometimes assumed) is a genus of an ornithopod dinosaur ...
'' and ''
Camptosaurus ''Camptosaurus'' ( ) is a genus of plant-eating, beaked ornithischian dinosaurs of the Late Jurassic period of western North America and possibly also Europe. The name means 'flexible lizard' (Greek (') meaning 'bent' and (') meaning 'lizard') ...
'', and the base is at about the same level as the
postzygapophyses The articular processes or zygapophyses (Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "process") of a vertebra are projections of the vertebra that serve the purpose of fitting with an adjacent vertebr ...
and
prezygapophyses The articular processes or zygapophyses (Greek language, Greek ζυγον = "yoke" (because it links two vertebrae) + απο = "away" + φυσις = "-physis, process") of a vertebra are projections of the vertebra that serve the purpose of fitting ...
. The prezygapophyses are just above the dorsal margin of the
neural canal In the developing chordate (including vertebrates), the neural tube is the embryonic precursor to the central nervous system, which is made up of the brain and spinal cord. The neural groove gradually deepens as the neural fold become elevated, ...
and are about 80 mm above the ventral rim of the centrum. The ribs are correlated with the possession of transverse processes that are inclined slightly upwards, a feature that differentiates it from ''
Dacentrurus ''Dacentrurus'' (meaning "tail full of points"), originally known as ''Omosaurus'', is a genus of stegosaurian dinosaur from the Late Jurassic to Early Cretaceous (154 - 140 mya) of Europe. Its type species, ''Omosaurus armatus'', was named in 18 ...
'', a stegosaur also from the Lourinhã Formation. The holotype preserves ossified tendons which is a feature seen in ankylosaurs,
ornithopods 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 possibly ''
Scelidosaurus ''Scelidosaurus'' (; with the intended meaning of "limb lizard", from Greek / meaning 'rib of beef' and ''sauros''/ meaning 'lizard')Liddell & Scott (1980). Greek-English Lexicon, Abridged Edition. Oxford University Press, Oxford, UK. is a gen ...
''. Suberbiola et al. (2005) noted that the manus of ''Dracopelta'' may have retained a primitive
phalangeal formula The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
, as in the nodosaurid ''
Sauropelta ''Sauropelta'' ( ; meaning 'lizard shield') is a genus of nodosaurid dinosaur that existed in the Early Cretaceous Period of North America. One species (''S. edwardsorum'') has been named although others may have existed. Anatomically, ''Saurop ...
'' and basal
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 body ...
n ''
Scutellosaurus ''Scutellosaurus'' ( ) is a genus of thyreophoran ornithischian dinosaur that lived approximately 196 million years ago during the early part of the Jurassic Period in what is now Arizona, USA. It is classified in Thyreophora, the armoured di ...
'', while ankylosaurids and
stegosaurs Stegosauria is a group of Herbivore, herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous Period (geology), periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what i ...
showed a reduced phalangeal formula. Suberbiola et al. suggested that derived ankylosaurids and stegosaurs convergently evolved a tridactyl pes, with the loss of digit I, while ''
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 ...
'' pertained a tetradactyl manus and a tridactyl pes. The authors interpreted that ''Dracopelta'' may have had
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often u ...
adaptations as the autopodial structure falls into the metapodial range of the basal thyreophorans ''Scutellosaurus'' and ''Scelidosaurus'', which were probably subcursorial. The small size of ''Dracopelta'' is also consistent with the interpretation.


Classification

Galton (1980) originally assigned ''Dracopelta'' to
Nodosauridae 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, wer ...
, based on the morphology of the osteoderms. However, Vickaryous et al. (2004) interpreted ''Dracopelta'' as
Ankylosauria 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. ...
''
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty ...
'' while Carpenter (2001) considered it as a ''
nomen dubium In binomial nomenclature, a ''nomen dubium'' (Latin for "doubtful name", plural ''nomina dubia'') is a scientific name that is of unknown or doubtful application. Zoology In case of a ''nomen dubium'' it may be impossible to determine whether a s ...
'' based on the lack off diagnostic characteristic of the holotype specimen and assigned it as
Polacanthidae Polacanthinae is a subfamily of ankylosaurs, most often nodosaurids, from the Late Jurassic through Early Cretaceous of Europe and potentially North America and Asia. The group is defined as the largest clade closer to ''Polacanthus foxii'' than ...
''incertae sedis'' based on the similarities between ''
Gargoyleosaurus ''Gargoyleosaurus'' (meaning "gargoyle lizard") is one of the earliest ankylosaurs known from reasonably complete fossil remains. The holotype was discovered in 1995 at the Bone Cabin Quarry West locality, in Albany County, Wyoming in exposures ...
'' and '' Gastonia'' in osteoderm morphology.Vickaryous, M. K., Maryanska, T., and Weishampel, D. B. (2004). Chapter Seventeen: Ankylosauria. in The Dinosauria (2nd edition), Weishampel, D. B., Dodson, P., and Osmólska, H., editors. University of California Press. Suberbiola et al. (2005) were unable to recognize any polacanthid or nodosaurid characteristics and considered ''Dracopelta'' as a valid taxon based on the presence of proximal
phalanges The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...
II and III as long as wide and distinctive osteoderm morphology. Due to the close proximities between the fauna of the
Morrison Formation The Morrison Formation is a distinctive sequence of Late Jurassic, Upper Jurassic sedimentary rock found in the western United States which has been the most fertile source of dinosaur fossils in North America. It is composed of mudstone, sandsto ...
and Lourinhã Formation, ''Dracopelta'' might share close affinities with ''Gargoyleosaurus'' or ''
Mymoorapelta ''Mymoorapelta'' (Meaning "Vannetta Moore and Pete and Marilyn Mygatt's shield" after a combination of the names of the discoverers of the Mygatt-Moore Quarry that fossils were originally collected from, and pɛltə "shield") is a monospecific ...
''.


Paleoenvironment

The holotype specimen of ''Dracopelta'' was obtained from a layer from the Assenta Member of the Lourinhã Formation, which dates to the uppermost lower and upper Tithonian stage of the Late Jurassic. The member represents a fluvial channel and preserves small coalified
plant Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
fragments. The member is characterised by a succession of fluvial sandstones insinuated by erosive surfaces showing fossilised roots and
bioturbation Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a pr ...
, indicating periodic subaerial exposure. ''Dracopelta'' would have coexisted with the
carcharodontosaurian Allosauroidea is a superfamily or clade of theropod dinosaurs which contains four families — the Metriacanthosauridae, Allosauridae, Carcharodontosauridae, and Neovenatoridae. Allosauroids, alongside the family Megalosauroidea, were among th ...
''
Lusovenator ''Lusovenator'' (meaning "Portuguese hunter") is a genus of carcharodontosaurian theropod dinosaur, from the Late Jurassic (Kimmeridgian) Praia de Amoreira Porto-Novo Member and the Late Jurassic-Early Cretaceous (Tithonian-Berriasian) Assenta Me ...
'', indeterminate
turiasauria Turiasauria is an unranked clade of basal sauropod dinosaurs known from Middle Jurassic to Early Cretaceous deposits in Europe, North America, and Africa. Description Turiasauria was originally erected by Royo-Torres et al. (2006) to include '' ...
n
sauropods 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
crocodylomorphs Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cr ...
, and an indeterminate ankylosaur, which may represent a second and more complete specimen of ''Dracopelta''.


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

{{Taxonbar, from=Q132761 Ankylosaurs Kimmeridgian life Late Jurassic dinosaurs of Europe Jurassic Portugal Fossils of Portugal Taxa named by Peter Galton Fossil taxa described in 1980