The sauropod hiatus is a period in the North American fossil record for most of the Late Cretaceous noted for its lack of sauropod remains. It may represent an extinction event, possibly caused by competition with ornithischian herbivores, habitat loss from the expansion of the Western Interior Seaway, or both. Alternatively, it has been argued that the hiatus represents a decrease in inland deposits that would have effectively preserved the animals, creating the illusion of an extinction. The sauropod hiatus ended shortly before the end of the Cretaceous, with the appearance of Alamosaurus , most likely an immigrant from South America, in the southern parts of North America.
Definite evidence of Late Cretaceous sauropods in North America was first discovered in 1922, when Charles Whitney Gilmore described Alamosaurus sanjuanensis. [1] The term "sauropod hiatus" was coined by researchers Spencer G. Lucas and Adrian P. Hunt in 1989 to describes how fossils of the clade become scarce in western North America near the beginning of the Late Cretaceous. [2] [3] Another gap in the North American sauropod record was noted in the Early Cretaceous, from the Berriasian to the Barremian, but this gap was partially filled by the discovery of fragmentary remains of a Camarasaurus -like sauropod from the Lakota Formation in 2014. The fossil record of sauropods in this part of the Early Cretaceous is poor globally, not only in North America. [4]
Around the end of the Early Jurassic, sauropods, a clade of large, long-necked dinosaurs, established themselves as the "dominant group" of large herbivores in land-based environments, possibly following a global warming event. [5] They achieved global distribution, becoming a significant and diverse component of Mesozoic ecosystems. Sauropods were highly successful in North America during the Late Jurassic; the Morrison Formation of the western United States preserves one of the best-known of all sauropod faunas, [6] with 24 recognized sauropod species. [7] The fossil record of North American sauropods is poor for the first half of the Early Cretaceous, [4] but later in the Early Cretaceous, several lineages of sauropod are known from North America, including turiasaurs such as Moabosaurus , brachiosaurids such as Cedarosaurus , and somphospondyls such as Astrophocaudia .
The North American sauropod hiatus began during the Cenomanian, the first age of the Late Cretaceous. [8] Several sauropods are known from the upper Albian or lower Cenomanian of North America. The brachiosaurid Abydosaurus [9] and indeterminate sauropod teeth [2] have been found in the Mussentuchit Member, with the maximum age of the Abydosaurus fossils calculated to be 104.46±0.95 Ma. [9] The age of the brachiosaurid Sonorasaurus is not well-constrained, but probably was between 98 and 110 Ma. [10] Sauropod bones have been reported from the Dakota Formation. [2] Remains possibly referable to Sauroposeidon have been found in strata with a maximal age calculated to be 104.4 Ma. [8]
The species epithet of Sauroposeidon proteles means "perfected before the end" in reference to its status as one of the last North American sauropods before the sauropod hiatus. [11]
The cause of the sauropod hiatus has been a subject of debate. Lucas and Hunt proposed two hypotheses: a lack of properly preserved remains or a genuine extinction followed by re-population from South America. [3] In the first scenario—rejected by Lucas and Hunt themselves but later further explained by Thomas Lehman in 2001—as sea levels rose, the sauropods became restricted to upland environments that were not as well represented in fossil record as coastal ones, before eventually migrating back to lowland areas as they became more suitable. [2] In the second, the North American sauropods became extinct, and those that appear near the end of the Cretaceous represent immigrants from South America via land bridges, belonging to a group known as titanosaurians. [3]
In 2011, Philip D. Mannion and Paul Upchurch re-evaluated the sauropod hiatus, taking numerous factors into account. They argued that the event represented sampling bias, as the sauropods' preferred inland habitat is not as well represented in the fossil record as coastal regions, also claiming that the existence of Cenomanian-aged Sonorasaurus thompsoni shortened the gap to the Turonian–early Campanian. [2]
However, in 2012, Michael D. D’Emic and Brady Z. Foreman countered the analysis of Mannion and Upchurch. They cited a recent cladistic analysis of Sauroposeidon , which found it to be a type of somphospondylan, as evidence that titanosaurians had not been present in North America before the Maastrichtian, supporting a recent migration, also questioning claims of earlier North American titanosaurs. They further explained issues with the dates of the previous study; for instance Sonorasaurus thompsoni, provided as an example of a sauropod present during the hiatus by Mannion and Upchurch, was originally named as either Albian or Cenomanian in age when described, contrary to claims by Mannion and Upchurch that it was Cenomanian. Even assuming that the dates of the 2011 study were correct, D'Emic and Foreman pointed out that the hiatus would still last from the mid-Cenomanian to the mid-Campanian. As for potential sampling bias, they explained how other groups of dinosaurs from the same areas as sauropods persist into the Late Cretaceous, while members of the clade itself disappear at around 98 Mya.
D'Emic and Foreman discussed multiple events that could have led to a regional extinction. In the late Albian, the Western Interior Seaway began to push further into North America. In contrast to an earlier argument by Mannion and Upchurch that this event took place before the sauropod hiatus, D'Emic and Foreman write that it occurred gradually over millions of years. According to them, this occurred either before or concurrently with the event. Alternatively, in a scenario doubted by Lucas and Hunt but taken into further consideration by D'Emic and Foreman, hadrosaurs (duck-billed dinosaurs) could have out-competed sauropods. While the former pair believed that the two groups' feeding methods were too different for this to occur, the latter researchers suggested that such competition could have been between animals at younger stages of growth. This scenario is also supported by the fossil record, as the last pre-hiatus sauropods are found alongside the first North American hadrosauroids. [8]
Whatever the cause of the sauropod hiatus, sauropods do not reappear in the North American fossil record until near the end of the Cretaceous. Various specimens have been claimed to represent Campanian records of sauropods in North America, [12] [13] but have since been re-dated as Maastrichtian [14] or reinterpreted as representing other groups of animals. [15] Some of these claimed records have been determined to be hadrosaur cervical vertebrae, which can superficially resemble titanosaur caudal vertebrae if poorly preserved and interpreted back-to-front. [15] Sauropods do not definitely reappear in the North American fossil record until the Maastrichtian, the last age of the Cretaceous, [15] although an unpublished study presented at the Society of Vertebrate Paleontology conference in 2018 reported the occurrence of a sauropod caudal vertebra in the Deadhorse Coulee Member of the Milk River Formation, dating to the Santonian. If confirmed, it would represent the northernmost occurrence of a sauropod in Late Cretaceous North America, as well as provide evidence against the "sauropod hiatus." [16] The post-hiatus sauropod records of North America are all titanosaurs, and are commonly considered to represent a single genus, Alamosaurus . Alamosaurus appears to have rapidly become the dominant large herbivore of southern North America. [17] Due to its large size, Alamosaurus is estimated to have contributed to the evolution of gigantism in North American Tyrannosauridae species such as Tyrannosaurus, [18] with whom it lived alongside.
Alamosaurus probably reached North America by crossing over from South America. [19] [20] [21] Other studies have viewed the idea that Alamosaurus could have crossed over from South America skeptically, as the only connection between the Americas at the time may have been island chains that titanosaurs would have been poorly suited to cross, and proposed that Alamosaurus reached North America from Asia instead. [2] However, this hypothesis is based on studies that did not include Alamosaurus's South American close relative Pellegrinisaurus . [20] Sauropods have been considered poorly adapted for high-latitude environments, which would have made the Bering land bridge between Asia and North America inhospitable for titanosaurs. [21] The first appearance of hadrosaurs in South America coincides with the first appearance of titanosaurs in North America, suggesting that titanosaurs may have dispersed north along the same routes as hadrosaurs dispersed south. [15] It has also been proposed that Alamosaurus belongs to a lineage of titanosaurs native to North America, [17] but no other known North American sauropod is closely related to Alamosaurus, rendering such a possibility unlikely. [15]
Though the "sauropod hiatus" originally referred to a North American absence of sauropods, [3] a similar trend has been noted in Europe, lasting from the late Coniacian to the Santonian. The European gap was discussed in a similar manner by Éric Buffetaut and Jean Le Loeuff in multiple 1990s studies. [2] Buffetaut and Loeuff suggested that this European hiatus reflected an extinction of sauropods in Europe, suggesting that new migrants eventually arrived from Africa. [2] Mannion and Upchurch could not prove an African birthplace for European titanosaurs, although they stated Southern Hemispheric ancestry for certain groups to be a possibility. [2] Due to the scarcity of fossil deposits from the European sauropod hiatus, Le Loeuff considered the existence of such a hiatus tentative. [22] Trackways and a sauropod tooth have been found in Europe during the supposed hiatus, suggesting that sauropods were present in Europe throughout the Late Cretaceous, though their fossils are very rare. [23] [24]
Sauroposeidon is a genus of sauropod dinosaur known from several incomplete specimens including a bone bed and fossilized trackways that have been found in the U.S. states of Oklahoma, Wyoming, and Texas.
Titanosaurs 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 thriving at the time of the extinction event at the end of the Cretaceous. This group includes some of the largest land animals known to have ever existed, such as Patagotitan—estimated at 37 m (121 ft) long with a weight of 69 tonnes —and the comparably-sized Argentinosaurus and Puertasaurus from the same region.
The Brachiosauridae are a family or clade of herbivorous, quadrupedal sauropod dinosaurs. Brachiosaurids had long necks that enabled them to access the leaves of tall trees that other sauropods would have been unable to reach. In addition, they possessed thick spoon-shaped teeth which helped them to consume tough plants more efficiently than other sauropods. They have also been characterized by a few unique traits or synapomorphies; dorsal vertebrae with 'rod-like' transverse processes and an ischium with an abbreviated pubic peduncle.
Alamosaurus is a genus of opisthocoelicaudiine titanosaurian sauropod dinosaurs containing a single known species, Alamosaurus sanjuanensis, from the Maastrichtian age of the Late Cretaceous period in what is now southwestern North America. Isolated vertebrae and limb bones indicate that it reached sizes comparable to Argentinosaurus and Puertasaurus, which would make it the absolute largest dinosaur known from North America. Its fossils have been recovered from a variety of rock formations spanning the Maastrichtian age. Specimens of a juvenile Alamosaurus sanjuanensis have been recovered from only a few meters below the Cretaceous-Paleogene boundary in Texas, making it among the last surviving non-avian dinosaur species. Alamosaurus is the only known sauropod to have inhabited North America after their nearly 30-million year absence from the North American fossil record and probably represents an immigrant from South America.
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. Fossil evidence suggests that macronarian dinosaurs lived from the Middle Jurassic (Bathonian) through the Late Cretaceous (Maastrichtian). Macronarians have been found globally, including discoveries in Argentina, the United States, Portugal, China, and Tanzania. Like other sauropods, 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 and Titanosauriformes, among several others. Titanosauriforms are particularly well known for being some of the largest terrestrial animals to ever exist.
Opisthocoelicaudia is a genus of sauropod dinosaur of the Late Cretaceous Period discovered in the Gobi Desert of Mongolia. The type species is Opisthocoelicaudia skarzynskii. A well-preserved skeleton lacking only the head and neck was unearthed in 1965 by Polish and Mongolian scientists, making Opisthocoelicaudia one of the best known sauropods from the Late Cretaceous. Tooth marks on this skeleton indicate that large carnivorous dinosaurs had fed on the carcass and possibly had carried away the now-missing parts. To date, only two additional, much less complete specimens are known, including part of a shoulder and a fragmentary tail. A relatively small sauropod, Opisthocoelicaudia measured about 11.4–13 m (37–43 ft) in length. Like other sauropods, it would have been characterised by a small head sitting on a very long neck and a barrel shaped trunk carried by four column-like legs. The name Opisthocoelicaudia means "posterior cavity tail", alluding to the unusual, opisthocoel condition of the anterior tail vertebrae that were concave on their posterior sides. This and other skeletal features lead researchers to propose that Opisthocoelicaudia was able to rear on its hindlegs.
Sonorasaurus is a genus of brachiosaurid dinosaur from the Early to Late Cretaceous. It was a herbivorous sauropod whose fossils have been found in southern Arizona in the United States. Its name, which means "Sonora lizard", comes from the Sonora River that flows in the Sonoran Desert where its fossils were first found. The type species is S. thompsoni, described by Ratkevich in 1998.
Gobititan is a genus of herbivorous sauropod dinosaur from the Aptian faunal stage of the Early Cretaceous. The name of this genus is derived from the Gobi desert region and the Titans of Greek mythology, which is a reference to its large body size. The specific name shenzhouensis, is derived from "Shenzhou", an ancient name for China.
Europasaurus is a basal macronarian sauropod, a form of quadrupedal herbivorous dinosaur. It lived during the Late Jurassic of northern Germany, and has been identified as an example of insular dwarfism resulting from the isolation of a sauropod population on an island within the Lower Saxony basin.
Saltasauridae is a family of armored herbivorous sauropods from the Upper Cretaceous. They are known from fossils found in South America, Africa, Asia, North America, and Europe. They are characterized by their vertebrae and feet, which are similar to those of Saltasaurus, the first of the group to be discovered and the source of the name. The last and largest of the group and only one found in North America, Alamosaurus, was thirty-four metres in length and one of the last sauropods to go extinct.
The Cloverly Formation is a geological formation of Early and Late Cretaceous age that is present in parts of Montana, Wyoming, Colorado and Utah in the western United States. It was named for a post office on the eastern side of the Bighorn Basin in Wyoming by N.H. Darton in 1904. The sedimentary rocks of formation were deposited in floodplain environments and contain vertebrate fossils, including a diverse assemblage of dinosaur remains. In 1973, the Cloverly Formation Site was designated as a National Natural Landmark by the National Park Service.
Lithostrotia is a clade of derived titanosaur sauropods that lived during the Early Cretaceous and Late Cretaceous. The group was defined by Upchurch et al. in 2004 as the most recent common ancestor of Malawisaurus and Saltasaurus and all the descendants of that ancestor. Lithostrotia is derived from the Ancient Greek lithostros, meaning "inlaid with stones", referring to the fact that many known lithostrotians are preserved with osteoderms. However, osteoderms are not a distinguishing feature of the group, as the two noted by Unchurch et al. include caudal vertebrae with strongly concave front faces (procoely), although the farthest vertebrae are not procoelous.
The Javelina Formation is a geological formation in Texas. Dating has shown that the strata date to the Maastrichtian stage of the Late Cretaceous, approximately 70 to 66.5 million years old. The middle part of the formation has been dated to about 69 million years ago plus or minus 1 million years and the top situated near the Cretaceous–Paleogene boundary, dated to 66 Ma ago. Dinosaur remains are among the fossils that have been recovered from the formation.
The Black Peaks Formation is a geological formation in Texas whose strata date back to the Late Cretaceous. Dinosaur remains and the pterosaur Quetzalcoatlus northropi have been among the fossils reported from the formation. The boundary with the underlying Javelina Formation has been estimated at about 66.5 million years old. The formation preserves the rays Rhombodus and Dasyatis, as well as many gar scales.
Xianshanosaurus is a genus of sauropod dinosaur from the Early Cretaceous (Aptian-Albian) of the Ruyang Basin in Henan Province, China. Its type and only species is Xianshanosaurus shijiagouensis. It was described in 2009 by a team of paleontologists led by Lü Junchang. Xianshanosaurus may be a titanosaur, and Daxiatitan may be its closest relative, but its evolutionary relationships remain controversial.
Paluxysuchus is an extinct genus of neosuchian crocodyliform known from the Early Cretaceous Twin Mountains Formation of north-central Texas. It contains a single species, Paluxysuchus newmani. Paluxysuchus is one of three crocodyliforms known from the Early Cretaceous of Texas, the others being Pachycheilosuchus and an unnamed species referred to as the "Glen Rose Form". Paluxysuchus has a long, flat skull that is probably transitional between the long and narrow skulls of many early neosuchians and the short and flat skulls of later neosuchians.
Savannasaurus is a genus of titanosaurian sauropod dinosaur from the Late Cretaceous Winton Formation of Queensland, Australia. It contains one species, Savannasaurus elliottorum, named in 2016 by Stephen Poropat and colleagues. The holotype and only known specimen, originally nicknamed "Wade", is the most complete specimen of an Australian sauropod, and is held at the Australian Age of Dinosaurs museum. Dinosaurs known from contemporary rocks include its close relative Diamantinasaurus and the theropod Australovenator; associated teeth suggest that Australovenator may have fed on the holotype specimen.
Dzharatitanis is a genus of sauropod from the Bissekty Formation in Uzbekistan, dating to the Turonian age of the Late Cretaceous. The genus contains a single species, Dzharatitanis kingi, named after geologist Christopher King, who contributed to the Cretaceous geology of Asia. It is currently one of two known sauropods from the Bissekty Formation, alongside an indeterminate titanosaur. In its original publication it was considered to be a member of Rebbachisauridae, but later papers considered it to be a titanosaur.
Teratopodus is an ichnogenus of titanosaurian sauropod footprint. It includes a single species, T. malarguensis, known from prints found in the Late Cretaceous Anacleto Formation of Argentina. The Teratopodus tracks represent some of the best sauropod pes tracks currently known from South America.