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''Brachytrachelopan'' is a short-necked
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 b ...
dinosaur Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
from the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
( Oxfordian to
Tithonian In the geological timescale, the Tithonian is the latest age (geology), age of the Late Jurassic Epoch and the uppermost stage (stratigraphy), stage of the Upper Jurassic Series. It spans the time between 149.2 ±0.7 annum, Ma and 143.1 ±0.6 (mi ...
) of
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
. The
holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
and only known specimen ( Museo Paleontológico Egidio Feruglio MPEF-PV 1716) was collected from an
erosion Erosion is the action of surface processes (such as Surface runoff, water flow or wind) that removes soil, Rock (geology), rock, or dissolved material from one location on the Earth's crust#Crust, Earth's crust and then sediment transport, tran ...
al exposure of
fluvial A river is a natural stream of fresh water that flows on land or inside caves towards another body of water at a lower elevation, such as an ocean, lake, or another river. A river may run dry before reaching the end of its course if it ru ...
sandstone Sandstone is a Clastic rock#Sedimentary clastic rocks, clastic sedimentary rock composed mainly of grain size, sand-sized (0.0625 to 2 mm) silicate mineral, silicate grains, Cementation (geology), cemented together by another mineral. Sand ...
within the
Cañadón Calcáreo Formation The Cañadón Calcáreo Formation is an Oxfordian (stage), Oxfordian to Kimmeridgian-aged geologic formation, from the Cañadón Asfalto Basin in Chubut Province, Argentina, a rift basin that started forming since the earliest Jurassic.Figari et ...
on a hill approximately north-northeast of Cerro Cóndor,
Chubut Province Chubut ( from Tehuelche language, Tehuelche 'transparent'; ) is a provinces of Argentina, province in southern Argentina, situated between the 42nd parallel south (the border with Río Negro Province), the 46th parallel south (bordering Santa ...
, in west-central
Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...
,
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...
. Though very incomplete, the skeletal elements recovered were found in articulation and include eight cervical, twelve
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 The fus ...
, and three sacral
vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...
e, as well as
proximal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...
portions of the posterior cervical
rib In vertebrate anatomy, ribs () are the long curved bones which form the rib cage, part of the axial skeleton. In most tetrapods, ribs surround the thoracic cavity, enabling the lungs to expand and thus facilitate breathing by expanding the ...
s and all the dorsal ribs, the
distal Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position provi ...
end of the left
femur The femur (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg. The Femo ...
, the proximal end of the left
tibia The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
, and the right ilium. Much of the specimen was probably lost to erosion many years before its discovery. The
type species In International_Code_of_Zoological_Nomenclature, 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 spe ...
is ''Brachytrachelopan mesai''. The
specific name Specific name may refer to: * in Database management systems, a system-assigned name that is unique within a particular database In taxonomy, either of these two meanings, each with its own set of rules: * Specific name (botany), the two-part (bino ...
honours Daniel Mesa, a local shepherd who discovered the specimen while searching for lost sheep. The
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
name translates as "short-necked Pan", Pan being the god of the shepherds.


Description

This
taxon In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
's very short neck (approximately 40% shorter than other
dicraeosaurid Dicraeosauridae is a family of diplodocoid sauropods who are the sister group to Diplodocidae. Dicraeosaurids are a part of the Flagellicaudata, along with Diplodocidae. Dicraeosauridae includes genera such as ''Amargasaurus'', ''Suuwassea'', '' ...
s and the shortest of any known sauropod) is evidence that this lineage specialized to fill an ecological niche not exploited by other members of this
infraorder Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...
. Small for a sauropod, ''Brachytrachelopan'' measured in length and in body mass. Rauhut ''et al.'' (2005, 670) note that the high degree of fusion present between the preserved neural arches and their respective centra, as well as fusion between the sacral centra, sacral neural arches, and sacral neural spines is evidence that the holotype does not represent a juvenile animal. Hence, the small body size is not a relic of
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the ovum, egg to adult. The term can also be used to refer to t ...
.


Distinguishing characteristics

Rauhut ''et al.'' (2005, 670) diagnose ''Brachytrachelopan'' as differing from all other sauropods in the following respects: "...individual cervical vertebrae being as long as, or shorter in anteroposterior length than, high posteriorly. Further
apomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to hav ...
...include a pronounced, pillar-like centropostzygapophyseal lamina in the cervical vertebrae, a pronounced anterior inclination in the mid-cervical neural spines, with the tip of the spine extending beyond the anterior end of the centrum, and anterior dorsal neural spines one to six with vertical bases and anteriorly flexed tips."


Classification

''Brachytrachelopan'' belongs to
Sauropoda Sauropoda (), whose members are known as sauropods (; from ''wikt:sauro-, sauro-'' + ''wikt:-pod, -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative t ...
and
Neosauropoda Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentina, Argentine paleontologist José Bonaparte and currently described as ''Saltasaurus loricatus'', ''Diplodocus longus'', and all animals directly descended from their most recent ...
from the group of
Diplodocoidea Diplodocoidea is a superfamily of sauropod dinosaurs, which included some of the longest animals of all time, including slender giants like ''Supersaurus'', ''Diplodocus'', ''Apatosaurus'', and ''Amphicoelias''. Most had very long necks and long, ...
and family
Dicraeosauridae Dicraeosauridae is a Family (biology), family of Diplodocoidea, diplodocoid sauropods who are the sister group to Diplodocidae. Dicraeosaurids are a part of the Flagellicaudata, along with Diplodocidae. Dicraeosauridae includes genera such as ''A ...
. Following a
cladistic analysis Cladistics ( ; from Ancient Greek 'branch') 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 ...
of 27 sauropod taxa and 154
anatomical Anatomy () is the branch of morphology concerned with the study of the internal structure of organisms and their parts. Anatomy is a branch of natural science that deals with the structural organization of living things. It is an old scien ...
characters, Rauhut ''et al.'' (2005, 671-672; Fig. 2) assigned ''Brachytrachelopan'' to the
Dicraeosauridae Dicraeosauridae is a Family (biology), family of Diplodocoidea, diplodocoid sauropods who are the sister group to Diplodocidae. Dicraeosaurids are a part of the Flagellicaudata, along with Diplodocidae. Dicraeosauridae includes genera such as ''A ...
, proposing that, within this
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
, it should be considered to have a
sister group 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 ...
relationship to the Late Jurassic
Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...
n taxon ''Dicraeosaurus,'' instead of to ''Amargasaurus'' from the
Lower Cretaceous Lower may refer to: * ''Lower'' (album), 2025 album by Benjamin Booker * Lower (surname) * Lower Township, New Jersey *Lower Receiver (firearms) * Lower Wick Gloucestershire, England See also * Nizhny {{Disambiguation ...
of South America. Rauhut et al. (2005, 671) conclude this is indicative of a rapid
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within ...
and dispersal of the Dicraeosauridae following the separation of the
continent A continent is any of several large geographical regions. Continents are generally identified by convention (norm), convention rather than any strict criteria. A continent could be a single large landmass, a part of a very large landmass, as ...
s of the Southern and
Northern Hemisphere The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
s during the latest
Middle Jurassic The Middle Jurassic is the second Epoch (geology), epoch of the Jurassic Period (geology), Period. It lasted from about 174.1 to 161.5 million years ago. Fossils of land-dwelling animals, such as dinosaurs, from the Middle Jurassic are relativel ...
. The following cladogram by Tschopp and colleagues (2015) shows the presumed relationships between members of the Dicraeosauridae:


Paleoecology

Rauhut and colleagues in 2005 noted that the tendency towards shorter-necks seen in dicraeosaurids, and most evident in ''Brachytrachelopan'', runs counter to the lengthening of the neck seen in most sauropod lineages (
brachiosaurid The Brachiosauridae ("arm lizards", from Greek ''brachion'' (βραχίων) = "arm" and ''sauros'' = "lizard") are a family or clade of herbivorous, quadrupedal sauropod dinosaurs. Brachiosaurids had long necks that enabled them to access the l ...
s,
titanosaur 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 ...
s,
diplodocid Diplodocids, or members of the family Diplodocidae ("double beams"), are a group of sauropod dinosaurs. The family includes some of the longest creatures ever to walk the Earth, including '' Diplodocus'' and '' Supersaurus'', some of which may ha ...
s, etc.) and indicates that this group of sauropods was "progressively adapting for low browsing and might have been specialized on specific food sources, as has been suggested for ''
Amargasaurus ''Amargasaurus'' (; "La Amarga lizard") is a genus of sauropod dinosaur from the Early Cretaceous epoch (geology), epoch (129.4–122.46 Mya (unit), mya) of what is now Argentina. The only known skeleton was discovered in 1984 and is virtually ...
'' and ''
Dicraeosaurus ''Dicraeosaurus'' (Gr. , ' "bifurcated, double-headed" + Gr. , ' "lizard") is a genus of diplodocoid sauropod dinosaur that lived in what is now Lindi Region, Tanzania during the late Jurassic period. The genus was named for the neural spines on ...
''." Moreover, the
morphology Morphology, from the Greek and meaning "study of shape", may refer to: Disciplines *Morphology (archaeology), study of the shapes or forms of artifacts *Morphology (astronomy), study of the shape of astronomical objects such as nebulae, galaxies, ...
of the cervical neural arches in ''Brachytrachelopan'' would have significantly restricted dorsal
flexion Motion, the process of movement, is described using specific anatomical terminology, anatomical terms. Motion includes movement of Organ (anatomy), organs, joints, Limb (anatomy), limbs, and specific sections of the body. The terminology used de ...
of the neck and most likely indicates that this sauropod was specialized to a diet of plants "growing at heights of between about 1 and 2 m." Rauhut and colleagues also suggested that diet may have been a limiting factor in body size among dicraeosaurids, and that this may have placed them in the same
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...
as "large low-browsing
iguanodontia Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
n
ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
s." Such large iguanodontians are absent from the Late Jurassic
Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
n
sediment Sediment is a solid material that is transported to a new location where it is deposited. It occurs naturally and, through the processes of weathering and erosion, is broken down and subsequently sediment transport, transported by the action of ...
s that have produced all known
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 preserve ...
s of dicraeosaurids, while they are abundant in similar
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s of the same age in
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, where dicraeosaurids are absent. This may indicate that large iguanodontians and dicraeosaurids (especially ''Brachytrachelopan'') were ecological analogs, resulting from
parallel evolution Parallel evolution is the similar development of a trait in distinct species that are not closely related, but share a similar original trait in response to similar evolutionary pressure.Zhang, J. and Kumar, S. 1997Detection of convergent and pa ...
in two distantly related dinosaurian lineages.


References


External links


National Geographic
{{Taxonbar, from=Q131670 Dicraeosauridae Dinosaur genera Late Jurassic dinosaurs Fossil taxa described in 2005 Dinosaurs of Argentina