Abditosaurus
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''Abditosaurus'' (meaning "forgotten lizard") is an extinct genus of
titanosauria Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thr ...
n
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 ...
dinosaur from 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'', the ...
(
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
)
Tremp Group The Tremp Formation ( es, Formación de Tremp, ca, Formació de Tremp), alternatively described as Tremp Group ( es, Grupo Tremp), is a geological formation in the comarca Pallars Jussà, Lleida, Spain. The formation is restricted to the Trem ...
of
Catalonia Catalonia (; ca, Catalunya ; Aranese Occitan: ''Catalonha'' ; es, Cataluña ) is an autonomous community of Spain, designated as a ''nationality'' by its Statute of Autonomy. Most of the territory (except the Val d'Aran) lies on the north ...
,
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
. The type and only species is ''Abditosaurus kuehnei''. Phylogenetic analyses recover it within a clade of South American and African saltasaurines, distinct from other
insular dwarf Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
sauropods from the European archipelago. ''Abditosaurus'' inhabited the Ibero-Armorican Island, a prehistoric island made up of what is now Spain, Portugal, and southern France, and would have been the largest titanosaur species in its environment.


History of study

While prospecting for fossil mammals in 1954, German paleontologist
Walter Georg Kühne Walter Georg Kühne (February 26, 1911 in Berlin – March 16, 1991 ibid.) was a German paleontologist, known as a "legendary explorer of Mesozoic mammals". He graduated in March 1930 from reform boarding school ''Schule am Meer'' on the island of ...
on 25 September discovered the remains of a large titanosaur at the Orcau-1 locality of the
Conques Formation The Tremp Formation ( es, Formación de Tremp, ca, Formació de Tremp), alternatively described as Tremp Group ( es, Grupo Tremp), is a geological Formation (geology), formation in the comarca Pallars Jussà, Lleida Province, Lleida, Spain. Th ...
of
Catalonia Catalonia (; ca, Catalunya ; Aranese Occitan: ''Catalonha'' ; es, Cataluña ) is an autonomous community of Spain, designated as a ''nationality'' by its Statute of Autonomy. Most of the territory (except the Val d'Aran) lies on the north ...
,
Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...
. It was the first major sauropod find in Spain. During the initial two weeks of excavation Kühne managed to identify around ten bones, some of which were packaged and sent to the Instituto Lucas Mallada (ILM) in
Madrid Madrid ( , ) is the capital and most populous city of Spain. The city has almost 3.4 million inhabitants and a metropolitan area population of approximately 6.7 million. It is the second-largest city in the European Union (EU), and ...
, while others remained at the site (jacketed for protection). Returning the following year, Kühne, alongside locals and researchers of the ILM, managed to identify ten more bones in addition to most of those left at the site. Like in 1954, some of the bones were transported to Madrid while the remainder were left at the site. These initial discoveries consisted of fragments of the ilium, parts 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 eithe ...
, dorsal vertebrae,
chevrons Chevron (often relating to V-shaped patterns) may refer to: Science and technology * Chevron (aerospace), sawtooth patterns on some jet engines * Chevron (anatomy), a bone * ''Eulithis testata'', a moth * Chevron (geology), a fold in rock lay ...
, 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 ...
, parts of the
femurs 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 ...
and a complete
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 roun ...
alongside undetermined bones. Although Kühne asked for additional excavations, funding had run out and work at the locality was stopped. In 1956, the fossil material recovered was assigned by
Albert-Félix de Lapparent Albert-Félix de Lapparent (1905–1975) was a French palaeontologist. He was also a Sulpician priest. He undertook a number of fossil-hunting explorations in the Sahara desert. He contributed greatly to our knowledge of dinosaurs and other prehist ...
and
Emiliano Aguirre Emiliano Aguirre Enríquez (5 October 1925 – 11 October 2021) was a Spanish paleontologist, known for his works at archaeological site of Atapuerca, whose excavations he directed from 1978 until his retirement in 1990. He received the Prince ...
to a new, but unnamed, species of the now dubious genus ''
Hypselosaurus ''Hypselosaurus'' (meaning 'highest lizard', from Greek meaning 'high' or 'lofty' and meaning 'lizard') is a dubious genus of titanosaurian sauropod that lived in southern France during the Late Cretaceous, approximately 70 million years ago in ...
''. It was not until 1984 and 1986 that additional expeditions to the area were conducted, helmed by Josep Vicenç Santafé of the Institut de Paleontologia de Sabadell,
Barcelona Barcelona ( , , ) is a city on the coast of northeastern Spain. It is the capital and largest city of the autonomous community of Catalonia, as well as the second most populous municipality of Spain. With a population of 1.6 million within ci ...
, and assisted by researchers of the Institut d’Estudis Ilerdencs,
Lleida Lleida (, ; Spanish: Lérida ) is a city in the west of Catalonia, Spain. It is the capital city of the province of Lleida. Geographically, it is located in the Catalan Central Depression. It is also the capital city of the Segrià comarca, as ...
. They identified a sternal plate and additional dorsal vertebrae. Another expedition was conducted from 2012 to 2014 by the Institut Català de Paleontologia, the
University of Zaragoza The University of Zaragoza, sometimes referred to as Saragossa University () is a public university with teaching campuses and research centres spread over the three provinces of Aragon, Spain. Founded in 1542, it is one of the oldest universiti ...
and the Museu de la Conca Dellà. This expedition recovered elements left at the site by the 1955 and 1986 digs, with additional material of the limbs, tooth fragments, ribs and an articulated series of dorsal and cervical vertebrae. Subsequent research managed to discover some of the historical material. In 2017, part of the initial finds were described. This finding was shown in the 2017 documentary "''Europe's Last Giant''". The
type specimen In biology, a type is a particular wiktionary:en:specimen, specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to a ...
, which has numerous accession numbers, is the most complete known European titanosaur skeleton, with the semi-articulated neck, preserving twelve of the estimated fourteen cervical vertebrae, being especially noteworthy. Complete and nearly complete sequences of neck vertebrae are very rare in sauropods; as of early 2022, only 27 sauropod specimens with complete or nearly complete necks were known.


Etymology

In 2022, the
type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...
''Abditosaurus kuehnei'' was named and described by Bernat Vila, Albert Sellés, Miguel Moreno-Azanza, Novella L. Razzolini, Alejandro Gil-Delgado, José Ignacio Canudo and Àngel Galobart. The generic name, ''Abditosaurus'', means "forgotten reptile", and is derived 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 ...
''abditus'', meaning "concealed," or "forgotten," as the fossil material was hidden for sixty years between the initial discovery and the last expedition that collected the last bones, and 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 ...
''sauros'', meaning "lizard". The specific name, ''kuehnei'', honors Walter Georg Kühne, the discoverer of the type specimen.


Description

''Abditosaurus'' is noted for its large size, and is the largest titanosaur from the Ibero-Armorican Island, a geological region composing most of Spain and Southern France. It is estimated to have reached a length of and mass of . The holotype includes an
ossified Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by Cell (biology), cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes ...
sternal rib and an ossified
calcaneum In humans and many other primates, the calcaneus (; from the Latin ''calcaneus'' or ''calcaneum'', meaning heel) or heel bone is a bone of the tarsus of the foot which constitutes the heel. In some other animals, it is the point of the hock. St ...
, elements which hitherto had not been unequivocally identified in titanosaur specimens, leading to speculation that they had fully disappeared in the group. These bones probably ossified due to the advanced individual age of the holotype, which was also proven by its bone structure.


Classification

The discoverers of ''Abditosaurus'' interpreted it as a member of
Saltasaurinae Saltasaurinae is a subfamily of titanosaurian sauropods known from the late Cretaceous period of South America, India and Madagascar. Description Saltasaurines are relatively small sauropods with the general body shape of a small head, long ne ...
, a clade of titanosaurs found in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
and
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
. Their phylogenetic analysis recovered the Egyptian sauropod ''
Paralititan ''Paralititan'' (meaning "tidal giant") was a giant titanosaurian sauropod dinosaur genus discovered in coastal deposits in the Upper Cretaceous Bahariya Formation of Egypt. It lived between 99.6 and 93.5 million years ago.Holtz, Thomas R. Jr. (2 ...
'' as the sister taxon of ''Abditosaurus'', with the Brazilian ''
Maxakalisaurus ''Maxakalisaurus'' is a genus of titanosaur dinosaur, found in the Adamantina Formation of Brazil, in the state of Minas Gerais in 1998. The genus name is derived from the tribe of the Maxakali; Topa is one of their divinities.Kellner, A.W.A., C ...
'' as their next-closest relative. The clade uniting these three species, in turn, formed a
sister clade In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to
Saltasaurini Saltasaurini is a tribe of titanosaur sauropods known from the Late Cretaceous of Patagonia, Argentina. The clade was named in 2007 by Leonardo Salgado and José Bonaparte for the "least inclusive clade comprising ''Neuquensaurus'' and ''Saltasau ...
. As such, ''Abditosaurus'' is not closely related to other European sauropods, the majority of which were recovered as opisthocoelicaudiines by their phylogenetic analysis.


Paleoecology

The only known specimen of ''Abditosaurus'' was found in the
Conques Formation The Tremp Formation ( es, Formación de Tremp, ca, Formació de Tremp), alternatively described as Tremp Group ( es, Grupo Tremp), is a geological Formation (geology), formation in the comarca Pallars Jussà, Lleida Province, Lleida, Spain. Th ...
, which dates to the early
Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval from ...
age of the
Cretaceous The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of th ...
. At this time, southern France, Spain, and Portugal formed an island, known as the Ibero-Armorican Island. Most titanosaur taxa of the Ibero-Armorican Island, such as ''
Lirainosaurus ''Lirainosaurus'' (meaning "slender lizard"; from the Basque ''lirain'', meaning "slender", and the Greek ''sauros'', meaning "lizard")is a genus of titanosaur sauropod which lived in what is now Spain. The type species, ''Lirainosaurus astibiae ...
'', ''
Atsinganosaurus ''Atsinganosaurus'' is a genus of titanosaurian sauropod dinosaur which existed in what is now France during the late Cretaceous period. Well-preserved remains (and the only known) of ''Atsinganosaurus'' were collected from the Grès à Reptiles ...
'', ''
Garrigatitan ''Garrigatitan'' (meaning "garrigue giant") is a genus of titanosaurian dinosaur from the late Cretaceous Period of the Grès à Reptiles Formation in France. The genus contains a single species, ''Garrigatitan meridionalis.'' Discovery and nami ...
'', ''
Ampelosaurus ''Ampelosaurus'' ( ; meaning "vine lizard") is a titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now France. Its type species is ''A. atacis'', named by Le Loeuff in 1995. A possible unnamed species has given ''Ampel ...
'', and '' Lohuecotitan'', were small or medium-sized. In contrast, ''Abditosaurus'' was significantly larger and lacked physiological adaptations to an island environment, suggesting that the species was a recent arrival in the ecosystem. It may be part of a broader trend of larger sauropods replacing smaller endemic species in Europe during the early Maastrichtian, as larger sauropods first appear in the Hațeg biota around the same time. ''Abditosaurus'' is associated with eggs of the
oospecies Egg fossils are the fossilized remains of eggs laid by ancient animals. As evidence of the physiological processes of an animal, egg fossils are considered a type of trace fossil. Under rare circumstances a fossil egg may preserve the remains of ...
'' Fusioolithus baghensis'', which closely resemble eggs found in Gondwana.


References

{{Taxonbar, from1=Q110852461, from2=Q110872931 Macronarians Fossils of Spain Fossil taxa described in 2022 Late Cretaceous dinosaurs of Europe Saltasaurids Sauropods of Europe Titanosaurs