Padillasaurus Leivaensis
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''Padillasaurus'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
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
titanosauriform Macronaria is a clade of sauropod dinosaurs. Macronarians are named after the large diameter of the nasal opening of their skull, known as the external naris, which exceeded the size of the orbit, the skull opening where the eye is located (hence ...
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 ...
known from the
Early Cretaceous The Early Cretaceous ( geochronological name) or the Lower Cretaceous (chronostratigraphic name), is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 145  Ma to 100.5 Ma. Geology Pro ...
(
Barremian The Barremian is an age in the geologic timescale (or a chronostratigraphic stage) between 129.4 ± 1.5 Ma (million years ago) and 121.4 ± 1.0 Ma). It is a subdivision of the Early Cretaceous Epoch (or Lower Cretaceous Series). It is preceded ...
stage)
Paja Formation Paja may refer to: * Paja, Iran, a village in Sari County * Paja (given name), a Serbian masculine name * Paja (surname) * Paja Formation, geologic formation in Colombia * Paja Brava :''Paja Brava is also used in Spanish speaking countries to ...
in
Colombia Colombia (, ; ), officially the Republic of Colombia, is a country in South America with insular regions in North America—near Nicaragua's Caribbean coast—as well as in the Pacific Ocean. The Colombian mainland is bordered by the Car ...
. It contains a single species, ''Padillasaurus leivaensis'', known only from a single partial
axial skeleton The axial skeleton is the part of the skeleton that consists of the bones of the head and trunk (anatomy), trunk of a vertebrate. In the human skeleton, it consists of 80 bones and is composed of six parts; the human skull, skull (22 bones), als ...
. Initially described as a brachiosaurid, it was considered to be the first
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 ...
n brachiosaurid ever discovered and named. Before its discovery, the only known brachiosaurid material on the continent was very fragmentary and from the
Jurassic The Jurassic ( ) is a Geological period, geologic period and System (stratigraphy), stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The J ...
period. However, a more recent study finds it to be a basal
somphospondylan Somphospondyli is an extinct clade of titanosauriform sauropods that lived from the Late Jurassic until the end of the Late Cretaceous, comprising all titanosauriforms more closely related to Titanosauria proper than Brachiosauridae. The remain ...
.


Discovery

The fossil was discovered by local farmers in a
limestone Limestone ( calcium carbonate ) is a type of carbonate sedimentary rock which is the main source of the material lime. It is composed mostly of the minerals calcite and aragonite, which are different crystal forms of . Limestone forms whe ...
nodule during the 1990s in Ricaurte, northeast of
Villa de Leyva Villa de Leyva, also called Villa de Leiva, is a touristic colonial town and municipality, in the Ricaurte Province, part of the Boyacá Department of Colombia. The town is a Colombian National Heritage Town and is on the tentative list for UNES ...
. The exact site of the find is unknown, however the rock matrix in which the fossil was located contains
ammonite 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 cuttlefish) ...
fossils bellowing to the species '' Gerhardtia galeatoides'' and ''
Lytoceras ''Lytoceras'' is an ammonite genus that was extant during most of the Jurassic and Cretaceous periods, and is the type genus for the family Lytoceratidae. These cephalopods were fast-moving nektonic carnivores. Description Shells of ''Lytoceras' ...
'' sp., that allow the establishment of their origin in the middle Paja Formation which has been placed in the late Barremian, around 130 million years old. The fossil consists of vertebrae from the torso, sacrum and tail: all presumed to be from one individual. This specimen is the holotype assigned to the catalogue number JACVM 0001; this includes a rear dorsal vertebra, a series from the last two sacral vertebrae and first eight caudal vertebrae (these caudal vertebrae were found without their
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 ...
). The sacral vertebrae have been identified as the fourth and fifth. This fossil was donated by the farmers and became part of the collection at the museum of the Junta de Acción Comunal de la Vereda de Monquirá. ''Padillasaurus'' was originally described and named by José L. Carballido, Diego Pol, Mary L. Parra Ruge, Santiago Padilla Bernal, María E. Páramo-Fonseca y Fernando Etayo-Serna in 2015. The name of the genus pays homage to Carlos Bernardo Padilla, the founder of the Centro de Investigaciones Paleontológicas in
Villa de Leyva Villa de Leyva, also called Villa de Leiva, is a touristic colonial town and municipality, in the Ricaurte Province, part of the Boyacá Department of Colombia. The town is a Colombian National Heritage Town and is on the tentative list for UNES ...
, Colombia. The specific name, ''leivaensis'', is named after Villa de Leyva, which is located nearby to where the
vertebra 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 ...
e of 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 were discovered. ''Padillasaurus'' is the first record of a Cretaceous brachiosaurid of South America; previous findings of that time were only known in North America. Although fragmentary remains of possible brachiosaurids are known from the Jurassic of
Patagonia Patagonia () refers to a geographical region that encompasses the southern end of South America, governed by Argentina and Chile. The region comprises the southern section of the Andes Mountains with lakes, fjords, temperate rainforests, and gl ...
in Argentina, ''Padillasaurus'' it is the first member officially named from South America. It is also the latest brachiosaurid known from the ancient continent of
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 ...
.


Description

''Padillasaurus'' was a medium-sized titanosauriform. Because of the few remains found is difficult to determine what would have been its body size, but may have had between 16 and 18 meters long and weigh about 10 tonnes. ''Padillasaurus'' is distinguished from other titanosauriforms in having its
caudal 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 ...
weakly expanded to the sides, and displaying divided side projections. The species shows some typical basic features of Titanosauriformes: the vertebra is opisthocoelic, with a convex front section and the back concave. The edges show large pleurocoels, pneumatic processes that connected to the hollow interior of the body of the vertebra, which consisted of several hollow cavities. The spine also had an additional lamina between the diapophysis and the body of the vertebra. The anterior caudal vertebrae are flattened or slightly concave from the front and from behind, or vice versa. The
neural arch 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 ...
es are positioned relatively forward. These have long ridges that run between the protuberances of the joint face and body of the vertebra. The side projections are oriented obliquely backwards and have a convex lower area. The caudal vertebrae have a height equal to half of its width.


Phylogeny

''Padillasaurus'' was initially classified in the family Brachiosauridae based on its morphology. It had cavities in the sacral vertebrae that had no connection with the hollow interior of the vertebra, therefore lacking a true
pleurocoel Skeletal pneumaticity is the presence of air spaces within bones. It is generally produced during development by excavation of bone by pneumatic diverticula (air sacs) from an air-filled space, such as the lungs or nasal cavity. Pneumatization is h ...
. This feature links it to the brachiosaurids. A
cladistic analysis Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups ("clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived chara ...
was unable to determine the exact relationships of ''Padillasaurus'' with other genera, finding 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 ...
among basal forms including ''Padillasaurus'', ''
Abydosaurus ''Abydosaurus'' (meaning " Abydos lizard") is a genus of brachiosaurid sauropod dinosaur known from skull and postcranial material found in upper Lower Cretaceous rocks of northeastern Utah, United States. Discovery ''Abydosaurus'' is one of th ...
'', ''
Brachiosaurus ''Brachiosaurus'' () is a genus of sauropod dinosaur that lived in North America during the Late Jurassic, about 154to 150million years ago. It was first described by American paleontologist Elmer S. Riggs in 1903 from fossils found in th ...
'' and ''
Giraffatitan ''Giraffatitan'' (name meaning "titanic giraffe") is a genus of sauropod dinosaur that lived during the late Jurassic Period (geology), Period (Kimmeridgian–Tithonian stages) in what is now Lindi Region, Tanzania. It was originally named as an ...
'', while ''
Cedarosaurus ''Cedarosaurus'' (meaning "Cedar lizard" - named after the Cedar Mountain Formation, in which it was discovered) was a nasal-crested macronarian dinosaur genus from the Early Cretaceous Period (Valanginian). It was a sauropod which lived in what i ...
'' and ''
Venenosaurus ''Venenosaurus'' ( ) was a sauropod dinosaur. The name literally means "poison lizard", and it was named so after the Poison Strip Member of the Cedar Mountain Formation in Utah, United States, where the fossils were discovered by a Denver Museum ...
'' were occasionally recovered as
sister taxa 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 ...
. Unlike Cretaceous brachiosaurids from North America, in which the spinous process of the caudal vertebrae leaned slightly forward, those of ''Padillasaurus'' were angled rearward.
Cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...
from Carballido ''et al''., 2015: In 2017, a study concluded that ''Padillasaurus'' was not a brachiosaurid but a basal member of the
Somphospondyli Somphospondyli is an extinct clade of titanosauriform sauropods that lived from the Late Jurassic until the end of the Late Cretaceous, comprising all titanosauriforms more closely related to Titanosauria proper than Brachiosauridae. The remains ...
.Philip D. Mannion, Ronan Allain & Olivier Moine, 2017, "The earliest known titanosauriform sauropod dinosaur and the evolution of Brachiosauridae", ''PeerJ'' 5: e3217


References

{{Taxonbar, from=Q20886620 Macronarians Early Cretaceous dinosaurs of South America Cretaceous Colombia Fossils of Colombia Paja Formation Fossil taxa described in 2015 Taxa named by María Páramo