Heleosaurus Temporal range: Early-Middle Permian, | |
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Drawing of the Holotype specimen of Heleosaurus scholtzi (from Carroll, 1976) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Synapsida |
Family: | † Varanopidae |
Subfamily: | † Mycterosaurinae |
Genus: | † Heleosaurus Broom, 1907 |
Type species | |
†Heleosaurus scholtzi Broom, 1907 |
Heleosaurus scholtzi is an extinct species of basal synapsids, known as pelycosaurs, in the family of Varanopidae during the middle Permian. [1] At first H. scholtzi was mistakenly classified as a diapsid. [2] Members of this family were carnivorous and had dermal armor, and somewhat resembled monitor lizards. [1] This family was the most geologically long lived, widespread, and diverse group of early amniotes. To date only two fossils have been found in the rocks of South Africa. [3] [4] One of these fossils is an aggregation of five individuals. [4] [5]
H. scholtzi was first described by Broom in 1907 who originally placed it as an early diapsid. [3] It is named for his student, T.J.R. Scholtz, and was originally called Galechirus scholtzi. [3] Later work placed it as an Eosuchian [6] in the family Younginidae [6] and even proposed as an ancestor for Archosauria. [6] More recent work has placed it where it is now within Mycterosaurinae in the family Varanopidae. [1] [5] The closely related Elliotsmithia longiceps has been placed in a sister taxon to H. scholtzi. [5]
Basal varanopids are small, slender animals specialized for a specific feeding niche. [1] H. scholtzi have six main features that are Varanopid autapomorphies, placing them firmly in the family. These are their slender, elongated quadratojugals; anterodorsal sloping of the quadrate; parasphenoid dentition; elongate hyoids; plate-like interclavicle heads; and their recurved and serrated teeth with a labiolingual compression. [1] They also have three main autapomorphies unique to the species; these are the trunk osteoderms, ornamentation on the surangular and angular, and a longitudinal median groove ventrally placed on the dorsal centra surface. [5]
The holotype fossil, SAM-PK-1070, is preserved in negative relief and has a partial skull, mandibles, axial skeleton, pectoral and pelvic girdles, osteoderms, and femur elements. The preserved femurs include a complete left and partial right. [1] [5] [3] Partial skull elements present are the maxilla, quadrate, parabasisphenol, jugal, quadratojugal, mandible, and the palate. Little of the skull roof remains in this specimen. The alveolar region of the mandible has five teeth preserved in it. [1]
The aggregate fossil composed of five individuals is better preserved and showed several new features not present in the holotype. These include a contact between the maxilla and both the prefrontal and the quadratojugals. [5] An anterior inclination of the occiput and the exclusion of the quadratojugal from the temporal fenestra are also clear in this specimen. [5] However, Spindler et al. (2018) transferred this aggregation to the separate species Microvaranops parentis. [7]
Many aspects of the skull structure are apparent between the two fossils. The overall shape is triangular and narrow, typical of Varanopids. [5] H. scholtzi’s snout is made up of the premaxilla bone. [5] A long narrow rectangular process of the naris along the antorbital region forms a straight dorsal border with the external naris. [5] Where the premaxilla meets with the naris there is a straight suture instead of the Varanopid typical V-shaped one. [5] The jugal and quadratojugal have pinched tubercular ornamentation on the lateral sides. [1] The quadratojugal has the elongated and slender shape distinct to the Varanopids, as well as a slender groove on the dorsal edge. This serves as an attachment for the squamosal, which extends anteriorly under the lateral temporal fenestra. [1] The right jugal is almost completely preserved with clear postorbital and subtemporal rami and a large tubercle near the ventral edge. [1] The vomers are long and slender with marginal teeth present on the medial edge of the right vomer. [1] H. scholtzi has ventrally preserved palantines that have extending diagonally across. Near the posterior edge of the internal naris they bifurcate. Suborbital fenestrae could not have been supported by the palantines due to their width. [1] Presence of this kind of fenestrae are seen in diapsids but absent in basal synapsids. [1] The parabasisphenoid has a cultriform process, the extended process at the anterior end of the braincase, that is transversely broad with a deep parabolic groove between cristae ventrolaterals. These are ridges formed by the lateral edges of the bones. Posteriorly paired depressions of the parabasisphenoid were likely for cervical muscle attachments. Small alveoli for teeth are also present on a continuous plate ventrally on the cultriform process. [1] Reisz (2007) describe a posterior coronoid, which was originally described as the prearticular by Carroll (1976). [1] [6] An anterior coronoid is also described by Reisz which was originally described as the splenial by Broom (1907). [6] [3] [1] The angular and surangular ornamentation are prolate and pustolous shaped tubercles [R]. Typical of Varanopids the ceratohyal is elongated and slender, acting like a strut that extends past the posterior end of the skull. [1] H. scholtzi’s parietals are broad, triangular, and take up most of the post-orbital region. The corners form a wing-like concave process postlaterally. The pineal foramen is large, oval in shape, and slightly sunken into the parietals. [5] Teeth of H. scholtzi are typical of Varanopids with a recurved shape, serrated mesially and distally on some. [1] A slightly larger caniniform is present a third of the way in from the anterior end of the maxilla. These teeth are very similar to those of Mycterosaurus, but a little more robust. [1]
Vertebrae of H. scholtzi have neural arches that are slightly longer anteroposteriorly than they are wide, a feature also seen in Mycterosaurus . [1] Neural spines have a blade like structure that are relatively narrow dorsally. [1] Most Varanopids have shallow grooves on the lateral surfaces of their vertebrae, but this is only seen in the cervical region of H. scholtzi. [1] This feature is also seen in Mesenosaurus. [1] The cervical vertebrae have a median ventral keel that is well developed. [1] Dorsal vertebrae have a broad ridge running along the midline of the centrum, subdivided by a shallow groove. This feature is unique and very uncommon in basal amniotes. [1] Articulation of the vertebrae can be seen in some of the individuals in the aggregate fossil, SAM-PK-K8305. This includes the caudal region of the vertebral column. [5] Slender ribs attached have holocephalous heads joined by a web of bones. [5] [1] The proximal ends are slender while the heads are triangular in shape. Very slender gastralia are present in the aggregate fossil. [1]
Surrounding the ventral column are a series of osteoderms, small rounded ossicles. [5] [1] These are a key autapomorphy of H. scholtzi and are articulated in the aggregate fossil. They form transverse rows of up to five osteoderms, with two to three rows per vertebrae. [5] This feature is shared by Elliotsmithia , another Permian synapsid. [8]
Preservation of the scapulacoracoids is good and show an autapomorphy for Varanopids, lack of a supraglenoid. [1] The scapular blade of H. scholtzi is low and broad. [1] Clavicles are preserved in both fossils and are long slender rods with narrow heads. [5] [1] These are positioned posterodorsally with the posterior edges tapering to connect with the scapulacoracoids. [5] The interclavicle head has a plate like shape with a long shaft, estimated by Reisz to be around the same length of six centra. [1]
The humerus is slender with narrow ends, well preserved in one individual of the aggregate fossil. [5] An ectepicondylar foramen is formed from the supinator process and ectiepicondyle. [5] Radii and ulna are preserved in the aggregate and are about equal in length, about 83% of the humerus length. The radius is almost straight and the ulna slightly more robust. [5] Two of the juvenile individuals of the aggregate have radii with a slightly bowed shape. [5] The metacarpals and phalanges of H. scholtzi have a long and slender shape to them. [5]
Pelvic girdles are preserved in both fossils and shows the anteroposteriorly elongated ischium typical of synapsids as well as a blade like distal shape and a well-developed pubic foramen. [1] The ilium is also elongated and rises anteriorly above the acetabulum. [5] At the anterodorsal margin of the acetabulum the ilium and pubis connect posteriorly to form it. [5] The pubis does not fuse to the ilium but instead twists to lay 90 degrees to the iliac blade. [5]
Femurs are preserved in both fossils and are slender, elongated bones with a sigmoidal curve. [9] [1] The proximal end turns up while the distal end curves down, forming the curved shape. [5] Shaft diameter of the femur is 10% of the total length and has a trochanter widely separated from the head. [1] Overall the femur is almost identical to that of Mycterosaurus. [10] [11] [1]
The exact locale of the holotype is uncertain due to Broom's inexact description, but work by Reisz and Modesto (2007) places it in the Tapinocephalus Assemblage Zone exposures in Victoria West. [1] This makes the horizon the Abrahamskraal formation, Beaufort Group, in the Karoo Basin. The discovery of the second fossil in that locale confirmed this as the horizon. [5]
Synapsida is a diverse group of tetrapod vertebrates that includes all mammals and their extinct relatives. It is one of the two major clades of the group Amniota, the other being the more diverse group Sauropsida. Unlike other amniotes, synapsids have a single temporal fenestra, an opening low in the skull roof behind each eye socket, leaving a bony arch beneath each; this accounts for the name "synapsid". The distinctive temporal fenestra developed about 318 million years ago during the Late Carboniferous period, when synapsids and sauropsids diverged, but was subsequently merged with the orbit in early mammals.
Eupelycosauria is a large clade of animals characterized by the unique shape of their skull, encompassing all mammals and their closest extinct relatives. They first appeared 308 million years ago during the Early Pennsylvanian epoch, with the fossils of Echinerpeton and perhaps an even earlier genus, Protoclepsydrops, representing just one of the many stages in the evolution of mammals, in contrast to their earlier amniote ancestors.
Petrolacosaurus is an extinct genus of diapsid reptile from the late Carboniferous period. It was a small, 40-centimetre (16 in) long reptile, and one of the earliest known reptile with two temporal fenestrae. This means that it was at the base of Diapsida, the largest and most successful radiation of reptiles that would eventually include all modern reptile groups, as well as dinosaurs and other famous extinct reptiles such as plesiosaurs, ichthyosaurs, and pterosaurs. However, Petrolacosaurus itself was part of Araeoscelida, a short-lived early branch of the diapsid family tree which went extinct in the mid-Permian.
Caseasauria is one of the two main clades of early synapsids, the other being the Eupelycosauria. Caseasaurs are currently known only from the Late Carboniferous and the Permian, and include two superficially different families, the small insectivorous or carnivorous Eothyrididae, and the large, herbivorous Caseidae. These two groups share a number of specialised features associated with the morphology of the snout and external naris.
Varanopidae is an extinct family of amniotes known from the Late Carboniferous to Middle Permian that resembled monitor lizards and may have filled a similar niche. Typically, they are considered to be relatively basal synapsids, although some studies from the late 2010s recovered them being taxonomically closer to diapsid reptiles, recent studies from the early 2020s support their traditional placement as synapsids on the basis of high degree of bone labyrinth ossification, maxillary canal morphology and phylogenetic analyses. A varanopid from the latest Middle Permian Pristerognathus Assemblage Zone is the youngest known varanopid and the last member of the "pelycosaur" group of synapsids.
Mycterosaurus is an extinct genus of amniotes belonging to the family Varanopidae. It is classified in the varanopid subfamily Mycterosaurinae. Mycterosaurus is the most primitive member of its family, existing from 290.1 to 272.5 MYA, known to Texas and Oklahoma. It lacks some features that its advanced relatives have.
Platyhystrix is an extinct temnospondyl amphibian with a distinctive sail along its back, similar to the unrelated synapsids, Dimetrodon and Edaphosaurus. It lived during the boundary between the latest Carboniferous and earliest Permian periods throughout what is now known as the Four Corners, Texas, and Kansas about 300 million years ago.
Eothyris is a genus of extinct synapsid in the family Eothyrididae from the early Permian. It was a carnivorous insectivorous animal, closely related to Oedaleops. Only the skull of Eothyris, first described in 1937, is known. It had a 6-centimetre-long (2.4-inch) skull, and its total estimated length was 30 centimetres. Eothyris is one of the most primitive synapsids known and is probably very similar to the common ancestor of all synapsids in many respects. The only known specimen of Eothyris was collected from the Artinskian-lower.
Angelosaurus is an extinct genus of herbivorous caseid synapsids that lived during the late Lower Permian (Kungurian) and early Middle Permian (Roadian) in what is now Texas and Oklahoma. Like other herbivorous caseids, it had a small head, large barrel-shaped body, long tail, and massive limbs. Angelosaurus differs from other caseids by the extreme massiveness of its bones, particularly those of the limbs, which show a strong development of ridges, processes, and rugosities for the attachment of muscles and tendons. Relative to its body size, the limbs of Angelosaurus were shorter and wider than those of other caseids. The ungual phalanges looked more like hooves than claws. The few known cranial elements show that the skull was short and more robust than that of the other representatives of the group. Angelosaurus is also distinguished by its bulbous teeth with shorter and wider crowns than those of other caseids. Their morphology and the high rate of wear they exhibit suggests a diet quite different from that of other large herbivorous caseids, and must have been based on particularly tough plants. A study published in 2022 suggests that the genus may be paraphyletic, with Angelosaurus possibly only represented by its type species A. dolani.
Dissorophus (DI-soh-ROH-fus) is an extinct genus of temnospondyl amphibian that lived during the Early Permian Period about 273 million years ago. Its fossils have been found in Texas and in Oklahoma in North America. Its heavy armor and robust build indicate Dissorophus was active on land, similar to other members of the clade Dissorophidae that are known from the Late Carboniferous to the Early Permian periods. Dissorphus is distinguished by its small body size, disproportionately large head and short trunk.
Aerosaurus is an extinct genus within Varanopidae, a family of non-mammalian synapsids. It lived between 252-299 million years ago during the Early Permian in North America. The name comes from Latin aes (aeris) “copper” and Greek sauros “lizard,” for El Cobre Canyon in northern New Mexico, where the type fossil was found and the site of former copper mines. Aerosaurus was a small to medium-bodied carnivorous synapsid characterized by its recurved teeth, triangular lateral temporal fenestra, and extended teeth row. Two species are recognized: A. greenleeorum (1937) and A. wellesi (1981).
Elliotsmithia is a small varanopid synapsid found from the late Middle Permian of South Africa. It is the sole basal synapsid "pelycosaur" known from the supercontinent Gondwana and only two specimens have been yielded to date. Its species name longiceps is derived from Latin, meaning "long head". Both known Elliotsmithia fossils were recovered from Abrahamskraal Formation rocks—within the boundaries of the Tapinocephalus Assemblage Zone—of the lower Beaufort Group. It was named for the late Sir Grafton Elliot Smith in 1937.
Mesenosaurus is an extinct genus of amniote. It belongs to the family Varanopidae. This genus includes two species: the type species Mesenosaurus romeri from the middle Permian Mezen River Basin of northern Russia, and Mesenosaurus efremovi from the early Permian (Artinskian) Richards Spur locality. M. romeri’s stratigraphic range is the middle to late Guadalupian while M. efremovi’s stratigraphic range is the Cisuralian.
Watongia is an extinct genus of non-mammalian synapsids from Middle Permian of Oklahoma. Only one species has been described, Watongia meieri, from the Chickasha Formation. It was assigned to family Gorgonopsidae by Olson and to Eotitanosuchia by Carroll. Reisz and collaborators assigned the genus in Varanopidae. Based on comparisons of its vertebrae with other varanopids, it was the largest varanopid with a body length of approximately 2 metres. It was a contemporary of its closest relative, the much smaller Varanodon; the two may possibly represent growth stages of a single animal.
Orovenator is an extinct genus of diapsid from Lower Permian deposits of Oklahoma, United States. It is known from two partial skulls from the Richards Spur locality in Oklahoma. The holotype OMNH 74606 consists of a partial skull preserving snout and mandible, and the referred specimen, OMNH 74607, a partial skull preserving the skull roof, vertebrae and palatal elements. It was first named by Robert R. Reisz, Sean P. Modesto and Diane M. Scott in 2011 and the type species is Orovenator mayorum. The generic name means "mountain", oro, in Greek in reference to the Richards Spur locality, which was mountainous during the Permian period and "hunter", venator, in Latin. The specific name honours Bill and Julie May. Orovenator is the oldest and most basal neodiapsid to date.
Ascendonanus is an extinct genus of varanopid amniote from the Early Permian of Germany. It is one of the earliest specialized arboreal (tree-living) tetrapods currently known and outwardly resembled a small lizard. The animal was about 40 cm long, with strongly curved claws, short limbs, a slender, elongated trunk, and a long tail. It would have preyed on insects and other small arthropods.
Dendromaia is an extinct genus of varanopid from the Carboniferous of Nova Scotia. It contains a single species, Dendromaia unamakiensis. Dendromaia is the oldest known varanopid, the only member of which to be discovered in Nova Scotia. Known from a large partial skeleton preserved with its tail wrapped around a much smaller partial skeleton, Dendromaia may also represent the oldest known occurrence of parental care in the fossil record. While the larger skeleton possessed certain mycterosaurine-like features, the smaller skeleton resembled basal varanopids such as Archaeovenator and Pyozia, creating uncertainty over whether characteristics at the base of Varanopidae have legitimate phylogenetic significance or instead reflect the immaturity of basal varanopid specimens.
Cabarzia is an extinct genus of varanopid from the Early Permian of Germany. It contains only a single species, Cabarzia trostheidei, which is based on a well-preserved skeleton found in red beds of the Goldlauter Formation. Cabarzia shared many similarities with Mesenosaurus romeri, although it did retain some differences, such as more curved claws, a wide ulnare, and muscle scars on its sacral ribs. With long, slender hindlimbs, a narrow body, an elongated tail, and short, thick forelimbs, Cabarzia was likely capable of running bipedally to escape from predators, a behavior shared by some modern lizards. It is the oldest animal known to have adaptations for bipedal locomotion, predating Eudibamus, a bipedal bolosaurid parareptile from the slightly younger Tambach Formation.
Anningia is an extinct genus in Varanopidae, a family of monitor lizard-like amniotes. It contains a single species, Anningia megalops.