Macrocnemus Temporal range: Middle Triassic, | |
---|---|
Various specimens of Macrocnemus from Monte San Giorgio in Switzerland. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Reptilia |
Clade: | Archosauromorpha |
Clade: | † Tanysauria |
Family: | † Tanystropheidae |
Genus: | † Macrocnemus Nopcsa, 1931 |
Species | |
|
Macrocnemus is an extinct genus of archosauromorph reptile known from the Middle Triassic (Late Anisian to Ladinian) of Europe and China. Macrocnemus is a member of the Tanystropheidae family and includes three species . Macrocnemus bassanii, the first species to be named and described, is known from the Besano Formation and adjacent paleontological sites in the Italian and Swiss Alps. [1] Macrocnemus fuyuanensis, on the other hand, is known from the Zhuganpo Formation in southern China. [2] A third species, Macrocnemus obristi, is known from the Prosanto Formation of Switzerland and is characterized by gracile limbs. [3] The name Macrocnemus is Greek for "long tibia".
Macrocnemus is known from multiple specimens, most belonging to M. bassanii. [4] It is a small reptile measuring 1.1–1.2 metres (3.6–3.9 ft) long. [5] Macrocnemus possessed at least 52 or 53 caudal vertebrae. Like many other early archosauromorphs, Macrocnemus had a small and low head on the end of a thin neck containing vertebrae with low neural spines and long cervical ribs. Many archosauromorphs with these features have been grouped within the order Protorosauria, although it is debatable whether this order is valid. [6] Features that are common to most "protorosaurs" like Macrocnemus include the ankle having a hooked fifth metatarsal attached to elongated limb elements, with tarsal elements with well-ossified proximal and distal ends. Unlike in Tanystropheus, digit V of the proximal phalanx of Macrocnemus is shorter than the other digits.
Macrocnemus bassanii is the most well-known and numerous species of Macrocnemus. Although the holotype specimen of this species was destroyed during World War II, a cast of the specimen (MSNM 14624, or alternatively PIMUZ T 2473) survived. [7]
Numerous complete or partial specimens of M. bassanni housed at PIMUZ (Paläontologisches Institut und Museum der Universität Zürich) include A III/208, T 1534, T 2470, T 2472, T 2474 through T 2477, T 2809, T 2812 through T 2816, T 4822, and T 4355. [4] [7]
The smallest specimen of this species is MSNM BES SC111, which is believed to be a juvenile about 30 centimeters long. The largest specimens were about 90 centimeters long.
Macrocnemus fuyuanensis was discovered at the Yunnan Province, southwestern China from the marine of the Triassic. More specially, it is known from the Zhuganpo Formation (formerly Zhuganpo Member of the Falang Formation). [8] This species in known from two nearly complete specimens, IVPP V15001 (the holotype specimen) and GMPKU-P-3001. The main features that set this species apart from M. bassanii include the presence of 17 or 18 dorsal vertebrae, a humerus which is longer than the radius, and a femur which is longer than the tibia. The front of the skull also differs compared to M. bassanii. According to Olivier Riepple et al, “a large, plate-like lacrimal is located in front of the tall, columnar prefrontal that defines the anterior margin of the orbit. A longitudinally oriented nasal groove extends along the anterior two-thirds of the snout, accommodating the external naris at its anterior part”. This species is also larger than M. bassanii. [2]
In 2017, a new specimen of Macrocnemus from the Besano Formation was described. This specimen, PIMUZ T 1559, possessed a long humerus, similar to that of M. fuyuanensis. It also differed from M. bassanii in the construction of its interclavicle. However, the interclavicles of the Chinese specimens are difficult to observe, and thus it is uncertain whether this European specimen is truly a member of M. fuyuanensis. Its describers placed it as "Macrocnemus aff. M. fuyuanensis", indicating its close relations to this species. [7]
Macrocnemus obristi was discovered by Christian Obrist during an excavation from the Upper Prosanto Formation, which is dated to the middle Triassic. It is known from two specimens, PIMUZ A/III 1467 (the holotype, consisting of a complete articulated tail, legs, and pelvic region) and PIMUZ A/III 722 (an isolated right tarsus). It is noticeably characterized by its gracile limb elements (including slender metatarsals) and a tibia which is 20% longer than the femur. Preserved soft tissue has also been found in the pelvic girdle of M. obristi's holotype. [3]
Some features of the limbs imply that Macrocnemus lived in terrestrial habitats and was capable of rapid bipedal movement. [4] Macrocnemus and Tanystropheus are among the most common "protorosaur" genera found in middle Triassic marine sediments. The presence of both Macrocnemus and Tanystropheus in both Switzerland and southwestern China suggest that the fauna of the western and eastern Tethyan realms was similar during the Middle and early Late Triassic. [2]
In 1970 Romer classified Macrocnemus as a lepidosaur related to modern lizards and tuataras, but in 1988 Carroll reclassified it as a member of Protorosauria. Other "protorosaurs" include Protorosaurus and Tanystropheus. Protorosaurs may not form a valid monophyletic clade according to some studies, which place tanystropheids as more derived archosauromorphs than other "protorosaurs". Macrocnemus is believed to be an early member of the tanystropheid family, as it shares many small skeletal features with Tanystropheus yet lacks the very elongated neck of that genus and other derived tanystropheids. [6]
Dinocephalosaurus is a genus of long necked, aquatic protorosaur that inhabited the Triassic seas of China. The genus contains the type and only known species, D. orientalis, which was named by Chun Li in 2003. Unlike other long-necked protorosaurs, Dinocephalosaurus convergently evolved a long neck not through elongation of individual neck vertebrae, but through the addition of neck vertebrae that each had a moderate length. As indicated by phylogenetic analyses, it belonged in a separate lineage that also included at least its closest relative Pectodens, which was named the Dinocephalosauridae in 2021. Like tanystropheids, however, Dinocephalosaurus probably used its long neck to hunt, utilizing the fang-like teeth of its jaws to ensnare prey; proposals that it employed suction feeding have not been universally accepted. It was probably a marine animal by necessity, as suggested by the poorly-ossified and paddle-like limbs which would have prevented it from going ashore.
Tanystropheus is an extinct genus of archosauromorph reptile which lived during the Triassic Period in Europe, Asia, and North America. It is recognisable by its extremely elongated neck, longer than the torso and tail combined. The neck was composed of 13 vertebrae strengthened by extensive cervical ribs. Tanystropheus is one of the most well-described non-archosauriform archosauromorphs, known from numerous fossils, including nearly complete skeletons. Some species within the genus may have reached a total length of 6 meters (20 ft), making Tanystropheus the longest non-archosauriform archosauromorph as well. Tanystropheus is the namesake of the family Tanystropheidae, a clade collecting many long-necked Triassic archosauromorphs previously described as "protorosaurs" or "prolacertiforms".
Mecistotrachelos is an extinct genus of gliding reptile from the Late Triassic of Virginia. It is generally interpreted as an archosauromorph, distantly related to crocodylians and dinosaurs. The type and only known species is M. apeoros. This specific name translates to "soaring longest neck", in reference to its gliding habits and long neck. This superficially lizard-like animal was able to spread its lengthened ribs and glide on wing-like membranes. Mecistotrachelos had a much longer neck than other gliding reptiles of the Triassic such as Icarosaurus and Kuehneosaurus. It was probably an arboreal insectivore.
Langobardisaurus is an extinct genus of tanystropheid archosauromorph reptile, with one valid species, L. pandolfii. Its fossils have been found in Italy and Austria, and it lived during the Late Triassic period, roughly 228 to 201 million years ago. Langobardisaurus was initially described in 1994, based on fossils from the Calcare di Zorzino Formation in Northern Italy. Fossils of the genus are also known from the Forni Dolostone of Northern Italy and the Seefeld Formation of Austria.
Sinosaurosphargis is an extinct genus of basal marine saurosphargid reptile known from the Middle Triassic Guanling Formation of Yunnan and Guizhou Provinces, southwestern China. It contains a single species, Sinosaurosphargis yunguiensis.
Cosesaurus is a genus of archosauromorph reptiles likely belonging to the family Tanystropheidae. It is known from fossil imprints of a single small skeleton, MGB V1, which was found in Muschelkalk outcrops near the municipalities of Mont-ral and Alcover in Spain. These outcrops are dated to the Ladinian age of the middle Triassic about 242 to 237 million years ago. The specimen is stored at the Museu Martorell, which is now part of the Museu de Ciències Naturals de Barcelona. The poor preservation and likely juvenile nature of the specimen has led to the anatomy of Cosesaurus being misidentified by several different sources. For example, Paul Ellenberger claimed that it was an ancestor to birds in the 1970s, while David Peters claimed that it was a pterosaur ancestor in 2000. Both of these claims contrast with mainstream scientific theories on the origins of either group, and other paleontologists who study the specimen are unable to find the features which Ellenberger or Peters reported to be present. The Ellenberger and Peters hypotheses are thus considered fringe theories with questionable scientific soundness due to their low reproducibility. Mainstream hypotheses for the relations of Cosesaurus generally agree that it is a "protorosaur", specifically a tanystropheid closely related to long-necked reptiles such as Macrocnemus, Tanytrachelos, Tanystropheus, or Langobardisaurus.
Protorosauria is an extinct, likely paraphyletic group of basal archosauromorph reptiles from the latest Middle Permian to the end of the Late Triassic of Asia, Europe and North America. It was named by the English anatomist and paleontologist Thomas Henry Huxley in 1871 as an order, originally to solely contain Protorosaurus. Other names which were once considered equivalent to Protorosauria include Prolacertiformes and Prolacertilia.
Pamelaria is an extinct genus of allokotosaurian archosauromorph reptile known from a single species, Pamelaria dolichotrachela, from the Middle Triassic of India. Pamelaria has sprawling legs, a long neck, and a pointed skull with nostrils positioned at the very tip of the snout. Among early archosauromorphs, Pamelaria is most similar to Prolacerta from the Early Triassic of South Africa and Antarctica. Both have been placed in the family Prolacertidae. Pamelaria, Prolacerta, and various other Permo-Triassic reptiles such as Protorosaurus and Tanystropheus have often been placed in a group of archosauromorphs called Protorosauria, which was regarded as one of the most basal group of archosauromorphs. However, more recent phylogenetic analyses indicate that Pamelaria and Prolacerta are more closely related to Archosauriformes than are Protorosaurus, Tanystropheus, and other protorosaurs, making Protorosauria a polyphyletic grouping.
Tanystropheidae is an extinct family of archosauromorph reptiles that lived throughout the Triassic Period, often considered to be "protorosaurs". They are characterized by their long, stiff necks formed from elongated cervical vertebrae with very long cervical ribs. Members of the group include both terrestrial and aquatic forms. While some tanystropheids were small lizard-like animals, other tanystropheids such as Tanystropheus were large animals that had necks that were several meters long, longer than the rest of their bodies.
Fuyuansaurus is an extinct genus of "protorosaur" reptiles known from the Middle Triassic Zhuganpo Formation of southern China. Fuyuansaurus was first named by Nicholas C. Fraser, Olivier Rieppel and Li Chun in 2013 and the type species is Fuyuansaurus acutirostris.
Eorasaurus is an extinct genus of archosauromorph reptile known from the middle late Permian of Tatarstan, European Russia. It contains a single species, Eorasaurus olsoni. When originally described by Sennikov (1997), Eorasaurus was identified as an early archosauromorph and assigned to the family Protorosauridae, Ezcurra et al. (2014) and Ezcurra (2016) later reclassified Eorasaurus and placed it within the group Archosauriformes. Eorasaurus is based solely on scant fossil material from the neck region, and is thus considered an unstable taxon in phylogenetic analyses. If Eorasaurus is an archosauriform, it would be the oldest known member of the group and would pre-date the previous record holder.
Helveticosauridae is an extinct family of basal marine reptiles known from the Middle Triassic of southern Switzerland and northern Italy.
Ozimek is a genus of sharovipterygid archosauromorph reptile, known from Late Triassic deposits in Poland and closely related to the Kyrgyzstani Sharovipteryx. It contains one species, O. volans, named in 2016 by Jerzy Dzik and Tomasz Sulej. Like Sharovipteryx, Ozimek had long, slender limbs with the hindlimbs longer than the forelimbs; the hindlimbs likely supported gliding membranes as fossilized in Sharovipteryx. Another unusual characteristic was the shoulder girdle, where the massive coracoids formed a shield-like structure covering the bottom of the shoulder region that would have limited mobility. In other respects, such as its long neck, it was a typical member of the non-natural grouping Protorosauria. Phylogenetic analysis has indicated that it, possibly along with Sharovipteryx, may have been an unusual member of the protorosaur group Tanystropheidae, although further study of its anatomy is needed to resolve its precise relationships.
Pectodens is an extinct genus of archosauromorph reptile which lived during the Middle Triassic in China. The type and only species of the genus is P. zhenyuensis, named by Chun Li and colleagues in 2017. It was a member of the Archosauromorpha, specifically part of the unnatural grouping Protorosauria. However, an unusual combination of traits similar and dissimilar to other protorosaurs initially led to confusion over its evolutionary relationships. In 2021, it was placed in a newly-established group, Dinocephalosauridae, along with its closest relative Dinocephalosaurus.
The Besano Formation is a geological formation in the southern Alps of northwestern Italy and southern Switzerland. This formation, a thin but fossiliferous succession of dolomite and black shale, is famous for its preservation of Middle Triassic (Anisian–Ladinian) marine life including fish and aquatic reptiles. It is exposed in the Monte San Giorgio and Besano area. It is among the formations responsible for the area being designated as a UNESCO World Heritage Site. In Switzerland, it is also known as the Grenzbitumenzone. The Anisian-Ladinian boundary lies in the upper part of the Besano Formation.
Elessaurus is an extinct genus of archosauromorph from the Early Triassic of Brazil. It contains a single species, Elessaurus gondwanoccidens. It possessed a variety of features common to basal archosauromorphs, particularly basal tanystropheids such as Macrocnemus. However, it is uncertain whether Elessaurus was a particularly close relative of tanystropheids, and it might instead be closer to other major archosauromorph clades. The genus name refers to "Elessar", an alternate name of the character Aragorn from J.R.R. Tolkien's Lord of the Rings trilogy.
Raibliania is an extinct genus of tanystropheid archosauromorph discovered in the Calcare del Predil Formation in Italy. It lived during the Carnian stage of the Late Triassic and it was related to Tanystropheus. Raibliania is distinct from Tanystropheus due to some distinct features of the cervical vertebrae and teeth. The type species is Raibliania calligarisi, named in 2020. The holotype consists of a partial post-cranial skeleton, with the known elements including vertebrae, a single tooth, several ribs, gastralia and parts of the pelvis.
The Zhuganpo Formation is a Triassic geologic unit found in southern China. It has historically been known as the Zhuganpo Member of the Falang Formation. A diverse fossil assemblage known as the Xingyi biota or Xingyi Fauna can be found in the upper part of the Zhuganpo Formation. Fossils of the Xingyi biota include articulated skeletons of marine reptiles, abundant fish, and a plentiful assortment of invertebrates indicating a Ladinian to Carnian age for the sediments of the formation.
Trachelosauridae is an extinct clade of archosauromorph reptiles that lived throughout the Triassic period. Like their close relatives the tanystropheids, they were "protorosaur"-grade archosauromorphs characterized by their long necks. Unlike tanystropheids, which lengthen their neck primarily by elongating the individual cervical (neck) vertebrae, trachelosaurids achieved their long necks by the addition of more vertebrae. The most extreme example of this trend was Dinocephalosaurus, which had at least 32 vertebrae in the neck alone, far more than the 13 neck vertebrae of Tanystropheus.
Gracilicollum is an extinct genus of likely tanysaurian archosauromorph from the Middle Triassic (Anisian) Guanling Formation of China. The genus contains a single species, G. latens, known from a skull and partial neck.