Aardonyx Temporal range: Early Jurassic, | |
---|---|
Reconstruction of the skull | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | † Sauropodomorpha |
Clade: | † Anchisauria |
Genus: | † Aardonyx Yates et al., 2010 |
Species: | †A. celestae |
Binomial name | |
†Aardonyx celestae Yates et al., 2010 | |
Aardonyx (Afrikaans aard, "earth" + Greek onux, "nail, claw") is a genus of basal sauropodomorph dinosaur. It is known from the type species Aardonyx celestae found from the Early Jurassic Elliot Formation of South Africa. A. celestae was named after Celeste Yates, who prepared much of the first known fossil material of the species. It has arm features that are intermediate between basal sauropodomorphs and more derived sauropods. [1]
Based on the structure of the hind limbs and pelvic girdle of Aardonyx, the dinosaur normally moved bipedally but could drop to quadrupedal movement similar to Iguanodon . It shares some attributes with giant quadrupedal sauropods like Apatosaurus . [2] Australian [3] [4] paleontologist Adam Yates and his team's discovery of the genus was published online before print in Proceedings of the Royal Society B in November 2009, and was scheduled to appear in the March 2010 issue. [2] British paleontologist Paul Barrett of the Natural History Museum, London, who was not involved in the research, commented that the discovery of Aardonyx "helps to fill a marked gap in our knowledge of sauropod evolution, showing how a primarily two-legged animal could start to acquire the specific features necessary for a life spent on all-fours". [2]
According to Matthew Bonnan, a co-author of the study, "We already knew that the earliest sauropods and near-sauropods would be bipeds. What Aardonyx shows us, however, is that walking quadrupedally and bearing weight on the inside of the foot is a trend that started very early in these dinosaurs, much earlier than previously hypothesized." Bonnan adds, "On a scientific level, it's really fulfilling to have a hypothesis on how you think dinosaurs got large, then to test that in the field and get back these kind of data — a new dinosaur — that really does start to fill in some of those anatomical gaps."
The genus is known from disarticulated bones belonging to two immature individuals. The material consists of cranial elements, vertebrae, dorsal and cervical ribs, gastralia, chevrons, elements of the pectoral and pelvic girdles, and bones of the fore and hind limbs, manus, and pes. The presence of these bones in a single dense accumulation in a localized channel fill suggests that they came from relatively complete carcasses. [5] Both individuals are thought to have been less than 10 years old at the time of their death because of the lack of peripheral rest lines in the cortices of sampled bones. Additional evidence for immaturity at the time of death includes calcified cartilage at the articular end of the scapula. [1]
Aardonyx is thought to be the sister taxon of a sauropodomorph clade containing Melanorosaurus and sauropods, which are all obligatory quadrupeds, based on a phylogenetic analysis conducted along with the first description of the genus. Many features of the skeleton support this relationship. These include derived traits seen in the vertebrae (such as hyposphenes that are as deep as the neural canal and mid-cervical neural spines that are less than twice as long as high) as well as the appendicular skeleton (such as the position of the fourth trochanter over the midlength of the femur and an adult femur length exceeding 600 mm). [1]
Cladogram showing the position of Aardonyx within Sauropodomorpha
after Yates et al., 2010: [1]
The following cladogram shows the position of Aardonyx within Massopoda, according to Oliver W. M. Rauhut and colleagues, 2020: [6]
Aardonyx shows a transition toward the bulk-browsing form of feeding characteristic of sauropods. The jaws of Aardonyx are narrow and V-shaped with a pointed symphysis, a plesiomorphic characteristic shared with other basal sauropodomorphs. In sauropods, the jaws are broad and U-shaped to allow for a wider bite. The absence of a lateral ridge at the caudal end of the dentary is indicative of a loss of fleshy cheeks. This is seen as an adaptation for a wider gape to facilitate bulk browsing and is observed in nearly all sauropods. The lateral neurovascular foramina of the maxilla of Aardonyx are smaller than those of other basal sauropodomorphs, and indicate that there was a reduction in blood supply to the buccal tissues and thus a loss of fleshy cheeks. The development of lateral plates along the alveolar margins of some bones of the skull would have helped brace the lingual sides of the teeth against bucco-lingual forces during foliage stripping.
The presence of plesiomorphic V-shaped jaws along with the absence of fleshy cheeks is an unusual characteristic of Aardonyx. Previously, it was thought that broader jaws evolved before the reduction and loss in fleshy cheeks as an adaptation toward bulk-browsing in sauropods. The sauropod Chinshakiangosaurus possessed jaws that were U-shaped, while still retaining fleshy cheeks, the opposite of the condition seen in Aardonyx. [7] Because Chinshakiangosaurus is a more derived sauropodomorph, this suggests that a wide, cheekless gape may have evolved twice in Sauropodomorpha: once with Aardonyx and again with sauropods more advanced than Chinshakiangosaurus. [1]
Characteristics of the limbs of Aardonyx suggest that it was habitually bipedal. Evidence for bipedalism can be seen in the forelimbs; the structure of the radius and ulna limited the degree to which the manus could be pronated, and the length of the humerus is only 72 percent that of the femur. However, characteristics found in both the fore and hind limbs of Aardonyx show a trend toward more habitual quadrupedalism that would eventually lead to the obligatory quadrupedalism seen in sauropods. The proximal end of the ulna possesses an incipient craniolateral process that gives the bone a y-shape similar to, although more subtle than, those of obligatory quadrupedal sauropodomorphs. The radius is shifted cranially, and a radial fossa allows for the ulna to cradle the radius craniolaterally. These characteristics suggest that there was a development towards greater quadrupedalism in Aardonyx. Although the hindlimbs of Aardonyx clearly show evidence for bipedalism (such as the retention of a convex proximal lateral profile of the femur and the position of the cranial trochanter far from the lateral margin of the femur), there is also evidence that indicates a shift toward quadrupedalism. Features of the femur suggests that the gait of Aardonyx was slower than that of more basal sauropodomorphs. The shaft of the femur is straighter and the fourth trochanter is more distally placed. The repositioning of the fourth trochanter to a more distal position causes the M. caudofemoralis longus muscle, the main femoral retractor muscle, to have greater leverage (more mechanical advantage) but conversely a decrease in the velocity of femoral retraction; consequently, Aardonyx was a powerful but slower walker than typical prosauropods.
Another characteristic that suggests a slower gait in Aardonyx is the robustness of metatarsal I in comparison with those of other basal sauropodomorphs. This is evidence of a more medial, or entaxonic, position of the weight-bearing axis of the foot, as opposed to a more mesaxonic position where the weight-bearing axis runs through digit III. The development of entaxony in Aardonyx provides further evidence for its reduced cursorial ability and wider gauge-gait, which is thought to have preceded obligatory quadrupedalism in sauropodomorphs. Previously, it was thought that entaxony developed after the divergence of Vulcanodon due to the presence of mesaxony in the genus. [8] However, the presence of mesaxony in Vulcanodon can be now considered an evolutionary reversal given the clear presence of entaxony in Aardonyx. [1]
The osteohistology of Aardonyx shows that it grew rapidly, but that its bodily growth was seasonally interrupted. Evidence for this comes from greatly vascularized woven-parallel complexes that contain regular growth marks, with its early and mid-ontogenetic stages exhibiting fibrolamellar complexes. [9]
Sauropoda, whose members are known as sauropods, is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads, and four thick, pillar-like legs. They are notable for the enormous sizes attained by some species, and the group includes the largest animals to have ever lived on land. Well-known genera include Apatosaurus, Argentinosaurus, Alamosaurus, Brachiosaurus, Camarasaurus, Diplodocus, and Mamenchisaurus.
Plateosaurus is a genus of plateosaurid dinosaur that lived during the Late Triassic period, around 214 to 204 million years ago, in what is now Central and Northern Europe. Plateosaurus is a basal (early) sauropodomorph dinosaur, a so-called "prosauropod". The type species is Plateosaurus trossingensis; before 2019, that honor was given to Plateosaurus engelhardti, but it was ruled as undiagnostic by the ICZN. Currently, there are three valid species; in addition to P. trossingensis, P. longiceps and P. gracilis are also known. However, others have been assigned in the past, and there is no broad consensus on the species taxonomy of plateosaurid dinosaurs. Similarly, there are a plethora of synonyms at the genus level.
Melanorosaurus is a genus of basal sauropodomorph dinosaur that lived during the Late Triassic period. An omnivore from South Africa, it had a large body and sturdy limbs, suggesting it moved quadrupedally. Its limb bones were massive and heavy like the limb bones of true sauropods.
Massospondylus was a genus of sauropodomorph dinosaur from the Early Jurassic. It was described by Sir Richard Owen in 1854 from remains discovered in South Africa, and is thus one of the first dinosaurs to have been named. Fossils have since been found at other locations in South Africa, Lesotho, and Zimbabwe. Material from Arizona's Kayenta Formation, India, and Argentina has been assigned to the genus at various times, but the Arizonan and Argentinian material are now assigned to other genera.
Isanosaurus is an extinct genus of sauropod dinosaur from Thailand. It was originally dated to approximately 210 million years ago during the Late Triassic, which would make it one of the oldest known sauropods. Its age was later considered uncertain, and may be Early Jurassic or even as young as Late Jurassic. The only species is Isanosaurus attavipachi. Though important for the understanding of sauropod origin and early evolution, Isanosaurus is poorly known. Exact relationships to other early sauropods remain unresolved.
Yunnanosaurus is an extinct genus of sauropodomorph dinosaur that lived approximately 199 to 183 million years ago in what is now the Yunnan Province, in China, for which it was named. Yunnanosaurus was a large sized, moderately-built, ground-dwelling, quadrupedal herbivore, that could also walk bipedally, and ranged in size from 7 meters (23 feet) long and 2 m (6.5 ft) high to 4 m (13 ft) high in the largest species.
Antetonitrus is a genus of sauropodiform dinosaur found in the Early Jurassic Elliot Formation of South Africa. The only species is Antetonitrus ingenipes. Sometimes considered a basal sauropod, it is crucial for the understanding of the origin and early evolution of this group. It was a quadrupedal herbivore, like its later relatives, but shows primitive adaptations to use the forelimbs for grasping, instead of purely for weight support.
Coloradisaurus is a genus of massospondylid sauropodomorph dinosaur. It lived during the Late Triassic period in what is now La Rioja Province, Argentina. It is known from two specimens collected from the Los Colorados Formation of the Ischigualasto-Villa Unión Basin.
Gryponyx is an extinct genus of massopod sauropodomorph known from southern Free State, central South Africa.
Chinshakiangosaurus is a genus of dinosaur and probably one of the most basal sauropods known. The only species, Chinshakiangosaurus chunghoensis, is known from a fragmentary skeleton found in Lower Jurassic rocks in China. Chinshakiangosaurus is one of the few basal sauropods with preserved skull bones and therefore important for the understanding of the early evolution of this group. It shows that early sauropods may have possessed fleshy cheeks.
Kotasaurus is a genus of sauropod dinosaur from the Early Jurassic period (Sinemurian–Pliensbachian). The only known species is Kotasaurus yamanpalliensis. It was discovered in the Kota Formation of Telangana, India and shared its habitat with the related Barapasaurus. So far the remains of at least 12 individuals are known. The greater part of the skeleton is known, but the skull is missing, with the exception of two teeth. Like some sauropods, it had a tail club that would have been used for intraspecific combat or interpspecific defense.
Massospondylidae is a family of early massopod dinosaurs that existed in Asia, Africa, North America, South America and Antarctica during the Late Triassic to the Early Jurassic periods. Several dinosaurs have been classified as massospondylids over the years. The largest cladistic analysis of early sauropodomorphs, which was presented by Apaldetti and colleagues in November 2011, found Adeopapposaurus, Coloradisaurus, Glacialisaurus, Massospondylus, Leyesaurus and Lufengosaurus to be massospondylids. This result supports many previous analyses that tested fewer taxa. However, this analysis found the two recently described North American massopods, Sarahsaurus and Seitaad, and the South African Ignavusaurus to nest outside Massospondylidae, as opposed to some provisional proposals. Earlier in 2011, Pradhania, a sauropodomorph from India, was tested for the first time in a large cladistic analysis and was found to be a relatively basal massospondylid. Mussaurus and Xixiposaurus may also be included within Massospondylidae.
Glacialisaurus is a genus of sauropodomorph dinosaur. It lived during the Pliensbachian stage of the Early Jurassic period around 186 to 182 million years ago in what is now the central region of the Transantarctic Mountains of Antarctica. It is known from two specimens; the holotype, a partial tarsus (ankle) and metatarsus, and a partial left femur. The fossils were collected by a team led by paleontologist William R. Hammer during a 1990–91 field expedition to the Hanson Formation of Antarctica. They were described in 2007, and made the basis of the new genus and species Glacialisaurus hammeri. The genus name translates as “icy” or "frozen lizard”, and the species name honors Hammer.
Matthew Bonnan is an American paleobiologist, a Professor of Biological Sciences at Stockton University, and as of 2021 a singer/songwriter. His research combines traditional descriptive and anatomical study with computer-aided morphometric analysis and modeling of vertebrate skeletons, and he is the co-discoverer of three new species of dinosaurs. He is the author of the book The Bare Bones: An Unconventional Evolutionary History of the Skeleton, designed to introduce undergraduates and curious lay readers to the anatomy and evolution of the vertebrate skeleton. Bonnan has a music/art outreach project, Once Upon Deep Time, a pop/rock song cycle about the evolution of hearing and our connection to the tree of life.
Eusauropoda is a derived clade of sauropod dinosaurs. Eusauropods represent the node-based group that includes all descendant sauropods starting with the basal eusauropods of Shunosaurus, and possibly Barapasaurus, and Amygdalodon, but excluding Vulcanodon and Rhoetosaurus. The Eusauropoda was coined in 1995 by Paul Upchurch to create a monophyletic new taxonomic group that would include all sauropods, except for the vulcanodontids.
Ignavusaurus is a genus of basal sauropodomorph dinosaur that lived during the Early Jurassic in what is now Lesotho. Its fossils were found in the Upper Elliot Formation which is probably Hettangian in age. It was described on the basis of a partial, well preserved articulated skeleton. The type species, I. rachelis, was described in 2010 by Spanish palaeontologist F. Knoll.
Sarahsaurus is a genus of basal sauropodomorph dinosaur which lived during the Early Jurassic period in what is now northeastern Arizona, United States.
Pulanesaura is an extinct genus of basal sauropodiform known from the Early Jurassic Upper Elliot Formation of the Free State, South Africa. It contains a single species, Pulanesaura eocollum, known from partial remains of at least two subadult to adult individuals.
Meroktenos is a genus of basal sauropodomorph dinosaur that lived during the Late Triassic period of what is now Lesotho.
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