Dicraeosauridae
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Dicraeosauridae is a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
diplodocoid 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, ...
sauropods who are the sister group to
Diplodocidae 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 hav ...
. Dicraeosaurids are a part of the
Flagellicaudata Flagellicaudata is a clade of Dinosauria. It belongs to Sauropoda and includes two families, the Dicraeosauridae and the Diplodocidae. Phylogeny The clade Flagellicaudata was erected by Harris and Dodson (2004) for the diplodocoid clade formed ...
, along with Diplodocidae. Dicraeosauridae includes genera such as ''
Amargasaurus ''Amargasaurus'' (; "La Amarga lizard") is a genus of sauropod dinosaur from the Early Cretaceous epoch (129.4–122.46 mya) of what is now Argentina. The only known skeleton was discovered in 1984 and is virtually complete, including a ...
'', ''
Suuwassea ''Suuwassea'' is a genus of Dicraeosauridae, dicraeosaurid sauropod dinosaur found in the Upper Jurassic strata of the Morrison Formation, located in southern Carbon County, Montana, Carbon County, Montana, United States. The fossil remains were ...
'', ''
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 ...
'', and ''
Brachytrachelopan ''Brachytrachelopan'' is a short-necked sauropod dinosaur from the Late Jurassic ( Oxfordian to Tithonian) of Argentina. The holotype and only known specimen (Museo Paleontológico Egidio Feruglio MPEF-PV 1716) was collected from an erosion ...
''. Specimens of this family have been found in North America, Asia, Africa, and South America. Their temporal range is from the Early or
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 ...
to 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 ...
. Few dicraeosaurids survived into the Cretaceous, the youngest of which was ''
Amargasaurus ''Amargasaurus'' (; "La Amarga lizard") is a genus of sauropod dinosaur from the Early Cretaceous epoch (129.4–122.46 mya) of what is now Argentina. The only known skeleton was discovered in 1984 and is virtually complete, including a ...
''. The group was first described by German paleontologist
Werner Janensch Werner Ernst Martin Janensch (11 November 1878 – 20 October 1969) was a German paleontologist and geologist. Biography Janensch was born at Herzberg (Elster). In addition to Friedrich von Huene, Janensch was probably Germany's most imp ...
in 1914 with the discovery of ''
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 ...
'' in Tanzania. Dicraeosauridae are distinct from other sauropods because of their relatively short neck size and small body size. The clade is
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
and well-supported phylogenetically with thirteen unambiguous
synapomorphies 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 have ...
uniting it. They diverged from Diplodocidae in the Mid-Jurassic, as evidenced by the diversity of dicraeosaurids in both South America and East Africa when
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 ...
was still united by land. However, there is some disagreement among paleontologists on the phylogenetic placement of ''
Suuwassea ''Suuwassea'' is a genus of Dicraeosauridae, dicraeosaurid sauropod dinosaur found in the Upper Jurassic strata of the Morrison Formation, located in southern Carbon County, Montana, Carbon County, Montana, United States. The fossil remains were ...
'', the only genus of the Dicraeosauridae to be found in North America. It has been characterized as a basal dicraeosaurid by some and a member of the Diplodocidae by others. The placement of ''Suuwassea'' within Dicraeosauridae or Diplodocidae has substantial biogeographic implications for the evolution of Dicraeosauridae.


Classification

Dicraeosaurids are a part of Diplodocoidea and are the sister group to Diploidocidae. In the past two decades, the known diversity of the group has doubled. However, the classification of ''Suuwassea'' as a dicraeosaurid is not universally agreed upon. Some phylogenetic analyses have found ''Suuwassea'' to be a basal diplodocoid instead of a dicraeosaurid. One 2015 analysis has even found ''
Dyslocosaurus ''Dyslocosaurus'' (meaning "hard-to-place lizard") is the name given in 1992 to a genus of sauropod dinosaur from the Late Jurassic Period of Wyoming, North America. The holotype or type specimen the genus is based on, AC 663, is part of the col ...
'' as a member of Dicraeosauridae. A 2016 reappraisal of ''
Amargatitanis ''Amargatitanis'' (meaning " Amarga giant") is a genus of dicraeosaurid sauropod dinosaur (a type of large, long-necked quadrupedal herbivorous dinosaur) from the Barremian-age (Lower Cretaceous) La Amarga Formation of Neuquén, Argentina. Cl ...
'' has placed it into the Dicraeosauridae, as well. In 2018 a new genus, '' Pilmatueia'', was described. Dicraeosaurids are differentiated from their sister group, diplodocids, and from most sauropods by their relatively small body size and short necks. Dicraeosaurids are advanced sauropods within the monophyletic clade
Neosauropoda Neosauropoda is a clade within Dinosauria, coined in 1986 by Argentine paleontologist José Bonaparte and currently described as ''Saltasaurus loricatus'', ''Diplodocus longus'', and all animals directly descended from their most recent common ...
, which is generally characterized by gigantism. The relatively small body size of dicraeosaurids make them an important outlier relative to other taxa in Neosauropoda.


Phylogeny

There have been several different proposed phylogenies of Dicraeosauridae and the intra-group cladistics are not resolved. ''Suuwassea'' is variably positioned as either a basal dicraeosaurid or a basal diplodocoid. The phylogeny published by Tschopp and colleagues in 2015 is as follows: Tschopp includes ''
Dyslocosaurus ''Dyslocosaurus'' (meaning "hard-to-place lizard") is the name given in 1992 to a genus of sauropod dinosaur from the Late Jurassic Period of Wyoming, North America. The holotype or type specimen the genus is based on, AC 663, is part of the col ...
'' and '' Dystrophaeus'' as dicraeosaurids, two genera traditionally not considered to be part of Dicraeosauridae. The specimens of ''Dystrophaeus viamelae'' are highly fragmentary, with only a few bones available for study including an ulna, partial scapula, partial dorsal vertebrae, a distal radius, and some metacarpals. ''Dyslocosaurus polyonychius'' also has extremely limited fossil evidence that only includes appendicular elements, and the position of it in Tschopp's phylogeny is therefore considered "preliminary". Several studies, however, do not include even ''Suuwassea'' in Dicraeosauridae, such as Sereno ''et al.'' (2007); and JD Harris (2006). Other studies, however, recover ''Suuwassea'' as a basal dicraeosaurid, including Whitlock (2010) and Salgado ''et al.'' (2006).


Paleobiology


Feeding behavior

As sauropods, dicraeosaurids are obligate herbivores. Due to their relatively small necks and skull shape, it has been deduced that dicraeosaurids and diplodocids primarily browsed close to the ground or at mid height. Among the dicraeosurids, only ''
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 ...
'' has well-preserved dentition. This makes it difficult for paleontologists to make definitive statements about Dicraeosauridae feeding behavior compared to diplodocid feeding behavior. However, compared to its known relatives, ''Dicraeosaurus'' is unique in that it has an equal number of teeth in the upper and lower jaw, though teeth in the lower jaw are replaced more slowly.


Anatomy

Dicraeosaurids are characterized by their relatively small body size, short necks, and long neural spines.Daniela Schwarz-Wings, 10th Annual Meeting of the European Association of Vertebrate Paleontologists. Royo-Torres, R, Gascó, F, and Alcalá, L., ''coord.'' ¡''Fundamental!'' 20: 1-290. 2012. They are 10–13 meters in body length. They share thirteen unambiguous
synapomorphies 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 have ...
including dorsal vertebrae without pleurocoels, the presence of a ventrally directed prong on the squamosal, and a subtriangular-shaped dentary symphysis.


Distribution and evolution

Dicraeosaurid specimens have been found in three continents - Africa, South America, and North America. The distribution of species is primarily Gondwonan, with the exception of the North American ''Suuwassea''. The presence of ''Suuwassea'' in North America is unique among dicraeosaurids, therefore making the proper taxonomic classification of ''Suuwassea'' essential. The group likely first diverged from the diplodocids in the middle Jurassic in North America and subsequently dispersed into Gondwana, with the most diversity in East Africa and South America. ''Amargasaurus'' was the latest surviving dicraeosaurid genus, living into the Early Cretaceous period.


Sources

{{Taxonbar, from=Q133812 Flagellicaudata Early Jurassic first appearances Early Cretaceous extinctions Prehistoric dinosaur families