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Macronaria is a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
of
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 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 ...
s. 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 ''macro''- meaning large, and –''naria'' meaning nose). Fossil evidence suggests that macronarian dinosaurs lived from the Middle
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 ...
(
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 ...
) through the Late
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 ...
(
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 ...
). Macronarians have been found globally, including discoveries in Argentina, the United States, Portugal, China, and Tanzania. Like other
sauropods 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 ...
, they are known to have inhabited primarily terrestrial areas, and little evidence exists to suggest that they spent much time in coastal environments. Macronarians are diagnosed through their distinct characters on their skulls, as well as appendicular and vertebral characters. Macronaria is composed of several subclades and families notably including
Camarasauridae Camarasauridae is a family of sauropod dinosaurs. Among sauropods, camarasaurids are small to medium-sized, with relatively short necks. They are visually identifiable by a short skull with large nares, and broad, spatulate teeth filling a thick ...
and
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 th ...
iformes, among several others. Titanosauriforms are particularly well known for being some of the largest terrestrial animals to ever exist. Macronaria was described by Wilson and Sereno who proposed the new subdivisions among the 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 ...
. Previously, this clade was thought to have ''
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
Camarasauridae Camarasauridae is a family of sauropod dinosaurs. Among sauropods, camarasaurids are small to medium-sized, with relatively short necks. They are visually identifiable by a short skull with large nares, and broad, spatulate teeth filling a thick ...
forming one sister group, and
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 th ...
oidea and
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, ...
forming another. This proposed shift with Macronaria placed Diplodocoidea as an outgroup to the new clade Macronaria, under which all other neosauropods would fall.


Anatomy

The following list includes some of the distinguishing characteristics that are diagnostic for Macronaria: * The long axis of the distal ischial shaft lie on the same plane * Middle and posterior
neural spines 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 ...
have distal ends that extend out transversely * Posterior neural spines extend at the tip forming a triangular process upwards. * The anterior chevrons of the tail have open proximal articulation * Robust, spatulate (spoon-like), and broad-crowned teeth * Crests formed by large protruding nasal * Elongate
metacarpals In human anatomy, the metacarpal bones or metacarpus form the intermediate part of the skeleton, skeletal hand located between the phalanges of the fingers and the carpal bones of the wrist, which forms the connection to the forearm. The metacarpa ...
* Forelimbs are relatively long in comparison to the hind limbs (most clearly demonstrated by brachiosaurs)


Paleobiology


Posture, body size, and locomotion

The posture of macronarians is characterized by a novel ‘wide-gauged’ locomotor style, particularly in
titanosaurs 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 ...
. '' ''While sauropods are known to be the giants of the dinosaurs, macronarians were not exclusively large-bodied. Macronarians show divergence in the evolution of body size with some members both increasing and decreasing in size from the primitive condition. For example, despite all being in the titanosaur clade, ''
Argentinosaurus ''Argentinosaurus'' is a genus of giant sauropod dinosaur that lived during the Late Cretaceous period in what is now Argentina. Although it is only known from fragmentary remains, ''Argentinosaurus'' is one of the largest known land animals o ...
'' reached enormous sizes (~50 tons), while ''
Saltasaurus ''Saltasaurus'' (which means "lizard from Salta") is a genus of saltasaurid dinosaur of the Late Cretaceous period of Argentina. Small among sauropods, though still heavy by the standards of modern creatures, ''Saltasaurus'' was characterized by ...
'' and ''
Magyarosaurus ''Magyarosaurus'' (" Magyar lizard") is a genus of dwarf sauropod dinosaur from late Cretaceous Period (early to late Maastrichtian) in Romania. It is one of the smallest-known adult sauropods, measuring only in length and in body mass. The t ...
'' reached only 1.5-3 metric tons, which is fairly small for sauropods. While some macronarians represent some of the largest terrestrial vertebrates ever known, other macronarians experienced a steady size decrease over time.


Biogeography

Sauropods, widely speaking, have been associated with both coastal and inland environments. It is believed that macronarians such as titanosaurs were strictly terrestrial and associated with inland environments such as
lacustrine A lake is an area filled with water, localized in a basin, surrounded by land, and distinct from any river or other outlet that serves to feed or drain the lake. Lakes lie on land and are not part of the ocean, although, like the much larger ...
systems. These findings are based on ‘wide-gauged’ trackways produced by titanosaurs which are strongly correlated with terrestrial sediments. It is also believed that macronarians have
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 ...
n origins. ''
Camarasaurus ''Camarasaurus'' ( ) was a genus of quadrupedal, herbivorous dinosaurs and is the most common North American sauropod fossil. Its fossil remains have been found in the Morrison Formation, dating to the Late Jurassic epoch (Kimmeridgian to Titho ...
'' is among the most commonly found dinosaur from the Late Jurassic deposits of the Morrison Formation in the US. Unlike ''Camarasaurus,'' Titanosaurs were most commonly found in the Southern Hemisphere with the exception of ''
Alamosaurus ''Alamosaurus'' (; meaning "Ojo Alamo lizard") is a genus of opisthocoelicaudiine titanosaurian sauropod dinosaurs, containing a single known species, ''Alamosaurus sanjuanensis'', from the late Cretaceous Period of what is now southern North Am ...
'' which was found in North America. There is strong geologic evidence that a land bridge between South and North America existed at the end of the Cretaceous allowing for dispersal of organisms between the two landmasses. Cretaceous sauropods are thought to have moved northward from South America, thus explaining the high density of South American sauropods, and the sole appearance of ''Alamosaurus'' in the Western Interior.


Diet

It is well established that all sauropods, including macronarians, were obligate herbivores. Unlike their sister group, the diplodocids, which were thought to feed on low-lying plants, camarasaurids and other macronarians likely had strongly upward-oriented necks for browsing trees and taller plants. Each tooth family in ''Camarasaurus'' is thought to have had a maximum of three replacement teeth and tooth formation took about 315 days. The replacement rate is thought to be one tooth every 62 days – this is about the same level or higher than non-sauropod herbivores, though it is lower than the replacement rates of the sister taxa, ''Diplodocus.'' It is thought that this may be related to the fact that the low-browsing taxa ingested more grit and thus needed to replace teeth more, while ''Camarasaurus'' and other macronarians fed on mid to upper canopy plants where exogenous grit is almost non-existent. Given the large body size of these neosauropod herbivores, it is thought that this type of niche partitioning, characterized by different species taking advantage of different resources, was necessary for coexistence. It was at one point believed that polished pebbles occasionally found with sauropod skeletons were
gastroliths A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In oth ...
. Gastroliths are stones intentionally swallowed to aid with digestion (as is seen in a variety of modern birds). However, more recent taphonomic and sedimentological evidence suggests that sauropods did not use stones for digestion due to the general rarity of finding gastric mill-like stones with sauropod remains, and low relative mass of the stones to the size of sauropod bodies. When gastrolith-like rocks are found with sauropods, it may be that they were accidentally ingested, or intentionally ingested for mineral uptake.


Phylogeny

Macronaria is nested within Neosauropoda and is sister to Diplodocoidea. These groups split around the mid- to late-Jurassic and end at the K/Pg boundary. In the less than 100 Myr that they lived, macronarians developed at least 216
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 ...
and
autapomorphies In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to t ...
. Despite the fact that macronarians are named after cranial openings, they are described by more appendicular synapomorphies. The species that survived late into the Cretaceous were characterized by stocky, wide-gauged posture, most notably the highly derived saltasaurines. Also under Macronaria are ''Camarasaurus'', ''
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
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 th ...
ia. Camarasauromorpha is the most basal group of Macronaria. The
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 ...
below follows José Luis Barco Rodríguez (2010). The cladogram below follows José L. Carballido, Oliver W. M. Rauhut, Diego Pol and Leonardo Salgado (2011). Simplified cladogram of Macronaria after D'Emic (2012). For alternative cladograms see also Mateus ''et al.'' (2011), Mannion ''et al.'' (2013).Mannion, P. D., Upchurch, P., Barnes, R. N., & Mateus, O. (2013). Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of the Linnean Society, 168(1), 98-206.


References

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