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''Spicomellus'' (meaning "collar of spikes") is an extinct genus of herbivorous ankylosaurian
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 ...
that lived in the supercontinent
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...
during the Middle Jurassic Period. The
type Type may refer to: Science and technology Computing * Typing, producing text via a keyboard, typewriter, etc. * Data type, collection of values used for computations. * File type * TYPE (DOS command), a command to display contents of a file. * Ty ...
and only known species is ''Spicomellus afer'', named and described in 2021. Its remains were found in the third subunit of the
El Mers Group The El Mers Group is a geological group in the Middle Atlas of Morocco. It is subdivided into 3 formations named the El Mers 1, 2 and 3 Formations respectively. It is a marine deposit primarily consisting of marl, with gypsum present in the uppe ...
(
Bathonian In the geologic timescale the Bathonian is an age and stage of the Middle Jurassic. It lasted from approximately 168.3 Ma to around 166.1 Ma (million years ago). The Bathonian Age succeeds the Bajocian Age and precedes the Callovian Age. Strat ...
- Callovian), near Boulahfa, south of
Boulemane Boulemane ( ber, ⴱⵓⵍⵎⴰⵏ; ar, بولمان) is a town in northern of Middle Atlas Mountains in Morocco. It is located at around , in the region of Fès-Meknès Fès-Meknès or Fez-Meknes ( ar, فاس-مكناس, fās maknās; be ...
, Fès-Meknès, Morocco. The genus name means "spiked collar", from the
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
'spica' meaning spike, and 'mellum' meaning spiked dog collar and the specific name 'afer' means "the African". During the Jurassic,
eurypoda 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 ...
n dinosaurs, in particular
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 ...
, were diverse and abundant in
Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pan ...
(nowadays the northern continents), but their remains are extremely rare in Gondwanan deposits, nowadays the southern continents. Nevertheless, the existence of fragmentary remains and trackways in the deposits of Gondwana indicates the presence of eurypodan taxa there. ''Spicomellus'' is the second described eurypodan taxon from
North Africa North Africa, or Northern Africa is a region encompassing the northern portion of the African continent. There is no singularly accepted scope for the region, and it is sometimes defined as stretching from the Atlantic shores of Mauritania in ...
, after ''
Adratiklit ''Adratiklit'' (meaning "mountain lizard") is an extinct genus of herbivorous stegosaurian dinosaur that lived on the supercontinent Gondwana during the Middle Jurassic period (168–164 million years ago). The genus contains a single species, '' ...
'', and the oldest known ankylosaur from anywhere in the world, with the possible exception of an unnamed
thyreophoran 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 ...
from the
Isle of Skye The Isle of Skye, or simply Skye (; gd, An t-Eilean Sgitheanach or ; sco, Isle o Skye), is the largest and northernmost of the major islands in the Inner Hebrides of Scotland. The island's peninsulas radiate from a mountainous hub dominated ...
,
Scotland Scotland (, ) is a country that is part of the United Kingdom. Covering the northern third of the island of Great Britain, mainland Scotland has a border with England to the southeast and is otherwise surrounded by the Atlantic Ocean to the ...
that could be up to 2 million years older than ''Spicomellus'', though it is still unknown if this older species was a
stegosaur Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
or an
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. ...
. The holotype, NHMUK PV R37412, is housed at the Natural History Museum in
London London is the capital and largest city of England and the United Kingdom, with a population of just under 9 million. It stands on the River Thames in south-east England at the head of a estuary down to the North Sea, and has been a majo ...
.


Discovery and naming

It is unknown when the holotype specimen, NHMUK PV R37412, was discovered, but what is known is that it was acquired by the Natural History Museum from a commercial fossil dealer, Moussa Direct, based in
Cambridge Cambridge ( ) is a university city and the county town in Cambridgeshire, England. It is located on the River Cam approximately north of London. As of the 2021 United Kingdom census, the population of Cambridge was 145,700. Cambridge bec ...
,
U.K. The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...
in 2019. Maidment initially believed that the fossil was a forgery, but after
CT scan A computed tomography scan (CT scan; formerly called computed axial tomography scan or CAT scan) is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers ...
ning the fossil, she drew the conclusion that it was a genuine fossil, and ''Spicomellus afer'' was described by Maidment et al., on the 23rd of September, 2021 in an article published online in paleontological journal ''
Nature Ecology & Evolution ''Nature Ecology and Evolution'' is an online-only monthly peer-reviewed scientific journal published by Nature Publishing Group covering all aspects of research on ecology and evolutionary biology. It was established in 2017. Its first and curre ...
''. 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 ...
of ''Spicomellus afer'' consists of a single
rib In vertebrate anatomy, ribs ( la, costae) are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the chest, enabling the lungs to expand and thus facilitate breathing by expanding the ches ...
with four co- ossified spines. This is a trait unique to ''Spicomellus'' and not known from any other
vertebrate 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, ...
. The holotype specimen was histologically sectioned to confirm that it was an ankylosaurian. The information about the locality of the fossil was provided by Direct and confirmed through discussion with the Moroccan fossil dealer who sold it to him. The locality was visited by S.C.R.M. and D.O. in 2019 and 2020, respectively, to study the
sedimentology Sedimentology encompasses the study of modern sediments such as sand, silt, and clay, and the processes that result in their formation (erosion and weathering), transport, deposition and diagenesis. Sedimentologists apply their understanding of mo ...
and
stratigraphy Stratigraphy is a branch of geology concerned with the study of rock (geology), rock layers (Stratum, strata) and layering (stratification). It is primarily used in the study of sedimentary rock, sedimentary and layered volcanic rocks. Stratigrap ...
of the area. They found that the formation consisted of shallow marine and continental mixed
clastic Clastic rocks are composed of fragments, or clasts, of pre-existing minerals and rock. A clast is a fragment of geological detritus,Essentials of Geology, 3rd Ed, Stephen Marshak, p. G-3 chunks, and smaller grains of rock broken off other rocks ...
, evaporitic and carbonate sediments.


Description

Despite being a basal ankylosaurian, the preserved dermal spikes of the holotype were fused to the bone, which is a trait unique to ''Spicomellus'' and not known from any other
vertebrate 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, ...
. Because the dermal spikes were fused to the bone instead of being attached to muscle tissue in all other
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. ...
s, this trait would probably have made it harder for the animal to move. Though its exact size is unknown, it was likely comparable in size to other
Middle Jurassic The Middle Jurassic is the second epoch of the Jurassic Period. It lasted from about 174.1 to 163.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relatively rare, but geological formations co ...
ankylosaurs like ''
Sarcolestes ''Sarcolestes'' (meaning "flesh robber") is an extinct genus of ankylosaurian ornithischian dinosaur from the Oxford Clay of England. The current type and only species is ''S. leedsi'', and the holotype is a single partial left mandible. The ...
'' and ''
Tianchiasaurus ''Tianchisaurus'' (meaning "heavenly pool lizard"), also invalidly called ''Tianchiasaurus'' and "Jurassosaurus", is a genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms ...
.'' This puts our best size estimates for ''Spicomellus'' at no longer than when fully grown.


Classification

At first, Susannah Maidment was unsure whether ''Spicomellus'' was a stegosaur or an ankylosaur, but Maidment ''et al.'' (2021) confirmed that ''Spicomellus'' is a basal ankylosaurian. It was probably closely related to the only other known ankylosaurs that were alive at the same time, ''
Sarcolestes ''Sarcolestes'' (meaning "flesh robber") is an extinct genus of ankylosaurian ornithischian dinosaur from the Oxford Clay of England. The current type and only species is ''S. leedsi'', and the holotype is a single partial left mandible. The ...
'' and ''
Tianchisaurus ''Tianchisaurus'' (meaning "heavenly pool lizard"), also invalidly called ''Tianchiasaurus'' and "Jurassosaurus", is a genus of ankylosaurian dinosaur from the Late Jurassic ( Oxfordian– Kimmeridgian)-aged Toutunhe Formation of China. I ...
.'' These species come from the U.K. and China, respectively.


Paleoecology

''Spicomellus'' is only known from the El Mers Group's El Mers III Formation. It coexisted with the
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 bo ...
''"Cetiosaurus" mogrebiensis'' (which may or may not be synonymous with the sauropod ''
Atlasaurus ''Atlasaurus'' is a genus of sauropod dinosaurs from Middle Jurassic (Bathonian to Callovian stages) beds in North Africa.M. Monbaron, D. A. Russell, and P. Taquet. (1999). ''Atlasaurus imelakei'' n.g., n.sp., a brachiosaurid-like sauropod from t ...
'' from the contemporaneous terrestrial
Guettioua Formation The Guettioua Formation is a geological formation in Morocco. It dates back to the Middle Jurassic.Weishampel, David B; et al. (2004). "Dinosaur distribution (Middle Jurassic, Africa)." In: Weishampel, David B.; Dodson, Peter; and Osmólska, Halszk ...
) and the
stegosaur Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
''
Adratiklit ''Adratiklit'' (meaning "mountain lizard") is an extinct genus of herbivorous stegosaurian dinosaur that lived on the supercontinent Gondwana during the Middle Jurassic period (168–164 million years ago). The genus contains a single species, '' ...
'', browsing on low-growing plants and roots and tubers. Predators of the ecosystem consist of indeterminate theropods (possible megalosaurids).J. Jenny, A. Le Marrec, and M. Monbaron. (1981). Les empreintes de pas de dinosauriens dans le Jurassique moyen du Haut Atlas central (Maroc): nouveaux gisements et precisions stratigraphiques. ''Geobos''. 14(3):427-431 The discovery of ''Spicomellus'' also shows that the two major thyreophoran groups ( Ankylosauria and
Stegosauria Stegosauria is a group of herbivorous ornithischian dinosaurs that lived during the Jurassic and early Cretaceous periods. Stegosaurian fossils have been found mostly in the Northern Hemisphere, predominantly in what is now North America, Europe, ...
) co-existed for over 20 million years, and implies that the extinction of the stegosaurs may have happened for other, still unknown reasons.


References

{{Taxonbar, from=Q108667860 Ankylosaurs Ornithischian genera Bathonian genera Middle Jurassic dinosaurs of Africa Jurassic Morocco Fossils of Morocco Fossil taxa described in 2021