Boii Temporal range: Carboniferous, | |
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The crushed skull of Boii, showing impressions (grey) and the underside of preserved bones (yellow) illustrated along with other tetrapod fragments by Frič (1883) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Order: | † Microsauria |
Family: | † Tuditanidae |
Genus: | † Boii Carroll, 1966 |
Type species | |
Boii crassidens Frič, 1876 | |
Synonyms | |
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Boii is an extinct genus of microsaur within the family Tuditanidae. [1] It was found in Carboniferous coal from mines near the community of Kounov in the Czech Republic. The only remains of the genus consist of a crushed skull, shoulder girdle bones, and scales, which were similar to microsaurian elements originally referred to Asaphestera . Boii can be characterized by its heavily sculptured skull, thin ventral plate of the clavicles, and a larger number of fangs on the roof of the mouth. [2] For many years the type and only known species, Boii crassidens, was considered to be a species of Sparodus , [3] until 1966 when Robert Carroll assigned it to its own genus. [4]
The specimen now designated as Boii crassidens was first described by Antonin Frič, one of the most notable paleontologists in the late 19th century region of Bohemia (now part of the Czech Republic). Frič made many contributions to knowledge of Carboniferous tetrapods during his lifetime, including a particular article published in "Sitzungsberichte der königlichen Böhmischen Gesellschaft der Wissenschaften in Prague", the most notable bohemian scientific journal of its day. The article, which was published in 1876 within volume 1875 of the journal, was a list of Carboniferous animals he and his associates recently discovered at gaskohle (bituminous coal) mines near the localities of Nýřany and Kounová. His list included short preliminary descriptions for many new genera and species of tetrapods, including Microbrachis, Branchiosaurus, Hyloplesion (at that time called Stelliosaurus), and Sparodus. [5]
Most of the "Saurier" (reptiles, amphibians, or other tetrapods) he discussed in the article hailed from the Nýřany mines, with only two diagnostic examples found within the Kounová mines. One of these was a supposed amphibian he designated as "Labyrinthodon" shwarzenbergii, which is now known to not be an amphibian at all, but rather an early synapsid named Macromerion . [6] The other specimen, which is now known as Boii crassidens, he named Batrachocephalus crassidens. This specimen consisted of a crushed skull, preserved on a slab and counterslab of coal. Jaw bones, scales, and shoulder girdle elements were preserved on the slabs as well. [5] It is currently held at the National Museum in Prague, with the designation ČGH 83. [4]
This article was only a preliminary review of the creatures Frič and his associates discovered. A more elaborate description was published in 1883 as part of a personal monograph focusing solely on the creatures discovered at these mines. This description renamed Batrachocephalus crassidens to Sparodus crassidens, as the genus name 'Batrachocephalus' was already taken by Batrachocephalus mino , an Indonesian species of catfish. Frič also assigned an isolated maxilla (upper jaw bone) with designation ČGH 124 to this species, although it was quite a bit larger than the maxilla of the crushed skull. [3] [4]
In a 1966 review of microsaurs published by Robert Carroll, "Sparodus" crassidens was not found to be a member of the genus Sparodus. Instead, it was re-evaluated as belonging to a family of early microsaurs known as the Tuditanidae. This prompted Carroll to create a new genus name for the specimens previously considered to belong to "Sparodus" crassidens. The new name he found was Boii crassidens, named after the Boii tribe which inhabited the area of Bohemia during the time of the Roman Republic. [4]
The crushed skull was pressed between two plates of coal which preserved the outer impressions of bones on both the underside and upper side of the specimen. Although not all of the bones were preserved, the outer impressions helped to reconstruct the structure of these missing bones. The impressions were used to reconstruct the skull and lower jaws, while the skull itself (which preserved the palate better than the impressions) was removed and encased in Canada balsam. The skull was initially believed to have been stout, approximately as long as it was wide. This 'frog-like' skull is responsible for the original genus name "Batrachocephalus", which is Greek for "frog head". [5] However, later reconstructions from Robert Carroll have interpreted this wide shape as a result of the crushing the skull experienced, with the skull actually being narrow and somewhat triangular, more similar in shape to that of a lizard rather than a frog. [2]
The orbits (eye holes) are roughly midway down the length of the skull. Unlike the fairly smooth skulls of most microsaurs, the skull of Boii is covered with numerous grooves and ridges which radiate from the middle of their respective bones. The maxillary and premaxillary bones forming the edge of the snout contained many conical teeth, about 30 per each side of the upper jaw. This number is somewhat larger than that of the eponymous tuditanid Tuditanus . These marginal (edge) teeth in general are slightly larger towards the front of the skull, although only to a small extent. [4]
The palate (roof of the mouth) also possesses teeth. The palatine bones, positioned right next to the maxillae, possess a fair number of teeth. The most notable assortment of palatal teeth are a bundle of large fangs preserved midway down the skull. Large palatal fangs are also shared by Sparodus, explaining how Boii crassidens was once considered to be part of that genus. Frič (1883) considered these fangs to have grown out of the vomers (from the front of the skull) or possibly the parasphenoid (from the back of the skull), [3] but Carroll (1966) reconstructed the fangs as being part of the long pterygoid bones, which were originally reported as being toothless. [4] [2] The dentaries (main bones of the lower jaws) are also preserved and covered with teeth similar to those of the upper jaws. A row of small pits run from the symphysis (chin) along the upper portion of the outer face of the bones. [4]
Some bones of the shoulder girdle are known. A pair of V-shaped bones are preserved behind the skull, which Frič (1883) identified as coracoids. However, research by Carroll has indicated that microsaurs did not possess coracoid bones, and that the bones identified by Frič were actually clavicles. Clavicles possess two blade-like regions pointing away from each other at right angles when seen from the front. The lower regions point inwards and laying along the chest and the upper regions point upwards along the sides of the body. The lower regions (a.k.a. ventral plates) of Boii's clavicles are very thin, akin to those of a reptile. [2] As typical for many tetrapods, Boii also possessed an interclavicle which was positioned behind the clavicles at the center of the chest. However, the only remnant of this bone in the specimen is a large yet indistinct impression. A single arm bone was also present on the specimen, although paleontologists disagree whether it was a humerus (according to Frič, 1883) [3] or a radius or ulna (according to Carroll, 1966). [4]
Boii possessed large scales, as preserved in the only known specimen. The scales of the back of the animal were tile-like with rounded corners. They tightly overlapped in alternating rows, and each possessed a bulging rear edge. The scales of the belly of the animal were also overlapping. However, these belly scales were wider than the back scales, and their rows were stacked so frequently that only a small portion of each individual scale is visible. [3]
Frič (1883) considered Boii crassidens to be a member of the family Branchiosauridae, along with other species of Sparodus. [3] Branchiosaurids, a group of small gilled temnospondyls, are now believed to be only distant relatives of Sparodus and Boii. In 1894, John William Dawson listed the three Sparodus species as microsaurs rather than branchiosaurids. He did not explicitly note each individual species (instead clumping them as "Sparodus sp."), because he was unsure whether they were all valid members of the same genus. [7]
Dawson's suspicions were rectified in 1966, when Carroll split Boii crassidens off of Sparodus. The skull structure of Boii was considered to be very similar to that Asaphestera (which itself was similar to Tuditanus), so he placed Boii along with Asaphestera in Tuditanidae. Tuditanids were basal microsaurs, which did not evolve the unusual adaptations of more advanced families of microsaurs. They were terrestrial, lizard-like creatures with well-developed legs and jaw joints set about as far back as the neck joint. Although Boii crassidens was quite old by microsaur standards, it is one of the last known species of tuditanids. [2] Carroll (1966) suggested that it may have been descended from Asaphestera, supposedly one of the earliest known microsaurs to have evolved. [4] However, as of 2020 Asaphestera has been recognized as a chimeric taxon, based on specimens of a potential eothyridid along with a newly-named microsaur, Steenerpeton . [8]
Adelospondyli is an order of elongated, presumably aquatic, Carboniferous amphibians. They have a robust skull roofed with solid bone, and orbits located towards the front of the skull. The limbs were almost certainly absent, although some historical sources reported them to be present. Despite the likely absence of limbs, adelospondyls retained a large part of the bony shoulder girdle. Adelospondyls have been assigned to a variety of groups in the past. They have traditionally been seen as members of the subclass Lepospondyli, related to other unusual early tetrapods such as "microsaurs", "nectrideans", and aïstopods. Analyses such as Ruta & Coates (2007) have offered an alternate classification scheme, arguing that adelospondyls were actually far removed from other lepospondyls, instead being stem-tetrapod stegocephalians closely related to the family Colosteidae.
Westlothiana is a genus of reptile-like tetrapod that lived about 338 million years ago during the latest part of the Viséan age of the Carboniferous. The genus is known from a single species, Westlothiana lizziae. It is the oldest known uncontroversial tetrapod, closely related to but not an amniote.
Eucritta is an extinct genus of stem-tetrapod from the Viséan epoch in the Carboniferous period of Scotland. The name of the type and only species, E. melanolimnetes is a homage to the 1954 horror film Creature from the Black Lagoon.
Cochleosaurus (“spoon lizard”, from the Latin cochlear "spoon" and Greek sauros “lizard”_ were medium-sized edopoid temnospondyls that lived in Euramerica during the Moscovian age. Two species, C. bohemicus and C. florensis, have been identified from the fossil record.
Diplovertebron is an extinct genus of embolomere that lived in the Late Carboniferous period (Moscovian), about 310 million years ago. Diplovertebron was a medium-sized animal, around 50 cm in length. Members of the genus inhabited European Carboniferous swamps in what is now the Czech Republic. They were closely related to larger swamp-dwelling tetrapods like Proterogyrinus and Anthracosaurus. However, Diplovertebron were much smaller than these large, crocodile-like creatures. Known from a single species, Diplovertebron punctatum, this genus has had a complicated history closely tied to Gephyrostegus, another genus of small, reptile-like amphibians.
Megalocephalus is an extinct genus of baphetid amphibian from the late Carboniferous of the British Isles and the United States (Ohio). It contains two species, M. pachycephalus and M. lineolatus.
Neopteroplax is an extinct genus of eogyrinid embolomere closely related to European genera such as Eogyrinus and Pteroplax. Members of this genus were among the largest embolomeres in North America. Neopteroplax is primarily known from a large skull found in Ohio, although fragmentary embolomere fossils from Texas and New Mexico have also been tentatively referred to the genus. Despite its similarities to specific European embolomeres, it can be distinguished from them due to a small number of skull and jaw features, most notably a lower surangular at the upper rear portion of the lower jaw.
Asaphestera is an extinct genus of a synapsid described on the basis of fossils from the Carboniferous of the Joggins locality in Nova Scotia, Canada. It was originally described as an undetermined lepospondyl and subsequently classified as a microsaur within the family Tuditanidae. A study published in May 2020 found that specimens referred to Asaphestera represented several unrelated species. Steen (1934)'s original species name Asaphestera platyris was retained for a skull which has been re-evaluated as the earliest known synapsid.
Acherontiscus is an extinct genus of stegocephalians that lived in the Early Carboniferous of Scotland. The type and only species is Acherontiscus caledoniae, named by paleontologist Robert Carroll in 1969. Members of this genus have an unusual combination of features which makes their placement within amphibian-grade tetrapods uncertain. They possess multi-bone vertebrae similar to those of embolomeres, but also a skull similar to lepospondyls. The only known specimen of Acherontiscus possessed an elongated body similar to that of a snake or eel. No limbs were preserved, and evidence for their presence in close relatives of Acherontiscus is dubious at best. Phylogenetic analyses created by Marcello Ruta and other paleontologists in the 2000s indicate that Acherontiscus is part of Adelospondyli, closely related to other snake-like animals such as Adelogyrinus and Dolichopareias. Adelospondyls are traditionally placed within the group Lepospondyli due to their fused vertebrae. Some analyses published since 2007 have argued that adelospondyls such as Acherontiscus may not actually be lepospondyls, instead being close relatives or members of the family Colosteidae. This would indicate that they evolved prior to the split between the tetrapod lineage that leads to reptiles (Reptiliomorpha) and the one that leads to modern amphibians (Batrachomorpha). Members of this genus were probably aquatic animals that were able to swim using snake-like movements.
Batropetes is an extinct genus of brachystelechid recumbirostran "microsaur". Batropetes lived during the Sakmarian stage[a] of the Early Permian. Fossils attributable to the type species B. fritschi have been collected from the town of Freital in Saxony, Germany, near the city of Dresden. Additional material has been found from the Saar-Nahe Basin in southwestern Germany and has been assigned to three additional species: B. niederkirchensis, B. palatinus, and B. appelensis.
Euryodus is an extinct genus of microsaur within the family Gymnarthridae. Euryodus is a Lepospondyl from the clade Microsauria that lived during the Lower Permian. The name comes from Greek, meaning ‘broad-tooth’. It has been found in the southern half of North America, from its original discovery in Texas up to Utah.
Odonterpeton is an extinct genus of "microsaur" from the Late Carboniferous of Ohio, containing the lone species Odonterpeton triangulare. It is known from a single partial skeleton preserving the skull, forelimbs, and the front part of the torso. The specimen was found in the abandoned Diamond Coal Mine of Linton, Ohio, a fossiliferous coal deposit dated to the late Moscovian stage, about 310 million years ago.
Trihecaton is an extinct genus of microsaur from the Late Pennsylvanian of Colorado. Known from a single species, Trihecaton howardinus, this genus is distinctive compared to other microsaurs due to possessing a number of plesiomorphic ("primitive") features relative to the rest of the group. These include large intercentra, folded enamel, and a large coronoid process of the jaw. Its classification is controversial due to combining a long body with strong limbs, features which typically are not present at the same time in other microsaurs. Due to its distinctiveness, Trihecaton has been given its own monospecific family, Trihecatontidae.
Scincosaurus is an extinct genus of nectridean tetrapodomorphs within the family Scincosauridae.
Ossinodus is an extinct genus of stem tetrapod. Fossils have been found from the Ducabrook Formation in Queensland, Australia dating back to the middle Visean stage of the Early Carboniferous (Mississippian). It was originally placed within the family Whatcheeriidae, but the absence of an intertemporal bone as suggested by a recent reconstruction of the skull based on fragmentary material may prove it to be stemward of all whatcheeriids.
Tuditanidae is an extinct family of microsaurian tetrapods. Fossils have been found from Nova Scotia, Ohio, and the Czech Republic and are Late Carboniferous in age.
Ymeria is an extinct genus of early stem tetrapod from the Devonian of Greenland. Of the two other genera of stem tetrapods from Greenland, Acanthostega and Ichthyostega, Ymeria is most closely related to Ichthyostega, though the single known specimen is smaller, the skull about 10 cm in length. A single interclavicle resembles that of Ichthyostega, an indication Ymeria may have resembled this genus in the post-cranial skeleton.
Altenglanerpeton is an extinct genus of microsaur tetrapod from the Late Carboniferous or Early Permian of Germany. Altenglanerpeton was named in 2012 after the Altenglan Formation in which it was found. The type and only species is A. schroederi.
Sigournea is a genus of stem tetrapod from the Early Carboniferous. The genus contains only one species, the type species Sigournea multidentata, which was named in 2006 by paleontologists John R. Bolt and R. Eric Lombard on the basis of a single lower jaw from Iowa. The jaw came from a fissure-fill deposit of the St. Louis Limestone that was exposed in a quarry near the town of Sigourney and dates to the Viséan stage, making it approximately 335 million years old. Bolt and Lombard named the genus after Sigourney. The species name multidentata alludes to the many teeth preserved in the jaw. The jaw, which is housed in the Field Museum and cataloged as FM PR 1820, curves strongly downward but was probably straight to begin with, having been deformed by the process of fossilization after the individual died. Rooted in the dentary bone along the outermost edge of the jaw are 88 small, pointed marginal teeth. An additional row of even smaller teeth runs along the coronoids, three bones positioned lengthwise along the lower boundary of the dentary on the inner surface of the lower jaw. Bolt and Lombard were able to classify Sigournea as an early member of Tetrapoda based on the presence of bone surfaces covered in pits and ridges, a single row of dentary teeth, a jaw joint that faces upward, and an open groove for a lateral line along the outer surface of the jaw, and on the absence of teeth on the prearticular bone or enlarged fangs on the coronoids. Sigournea differs from other stem tetrapods in having several holes within a depression called the exomeckelian fenestra on the inner surface of the jaw.
Andersonerpeton is an extinct genus of aïstopod from the Bashkirian of Nova Scotia, Canada. It is known from a single jaw, which shares an unusual combination of features from both other aistopods and from stem-tetrapod tetrapodomorph fish. As a result, Andersonerpeton is significant for supporting a new classification scheme which states that aistopods evolved much earlier than previously expected. The genus contains a single species, A. longidentatum, which was previously believed to have been a species of the microsaur Hylerpeton.