Spathicephalus Temporal range: Early - Late Carboniferous, | |
---|---|
Life restoration of Spathicephalus mirus | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Sarcopterygii |
Clade: | Tetrapodomorpha |
Clade: | Elpistostegalia |
Clade: | Stegocephali |
Superfamily: | † Baphetoidea |
Family: | † Spathicephalidae Beaumont, 1977 |
Genus: | † Spathicephalus Watson, 1929 |
Species | |
Spathicephalus is an extinct genus of stem tetrapods (early four-limbed vertebrates) that lived during the middle of the Carboniferous Period. The genus includes two species: the type species S. mirus from Scotland, which is known from two mostly complete skulls and other cranial material, and the species S. pereger from Nova Scotia, which is known from a single fragment of the skull table. Based on the S. mirus material, the appearance of Spathicephalus is unlike that of any other early tetrapod, with a flattened, square-shaped skull and jaws lined with hundreds of very small chisel-like teeth. However, Spathicephalus shares several anatomical features with a family of stem tetrapods called Baphetidae, leading most paleontologists who have studied the genus to place it within a larger group called Baphetoidea, often as part of its own monotypic family Spathicephalidae. Spathicephalus is thought to have fed on aquatic invertebrates through a combination of suction feeding and filter feeding.
The type species of Spathicephalus, S. mirus, was named by paleontologist D. M. S. Watson in 1929. Watson described seven fossil specimens from an outcrop of the Rumbles Ironstone in the town of Loanhead in Midlothian, Scotland. The ironstone dates to the late Namurian stage (earliest Upper Carboniferous) and is part of the Limestone Coal Group. These specimens were discovered in the 1880s and include a mostly complete skull with the palate exposed, an impression of the underside of a skull roof, a right portion of the back of a skull, and various jaw fragments. [2] At the time, Spathicephalus and other tetrapods from the Namurian of Scotland were some of the oldest tetrapods known, predating the better-known Late Carboniferous tetrapod assemblages of the British Coal Measures. In November 1974, Scottish paleontologist Stanley P. Wood discovered additional skull and jaw fragments of Spathicephalus in an open-pit mine (the Dora Open Cast Mine) near the town of Cowdenbeath in Fife. Wood found these fossils in a layer of siltstone that is the same age as the ironstone in Loanhead. [3]
American paleontologist Donald Baird named a second species of Spathicephalus, S. pereger, from Nova Scotia in 1962. [2] Baird named S. pereger on the basis of an impression of the right half of a skull table that collectors from the Harvard Museum of Comparative Zoology found on a beach between Point Edward and Keating Cove on Cape Breton Island. The impression was preserved in siltstone from the Point Edward Formation, which dates to the latest Upper Carboniferous (equivalent to the early Namurian in Europe), meaning that S. pereger predates S. mirus by several million years. [4]
Unlike most other early tetrapods which have rounded or pointed snouts, Spathicephalus mirus has a flattened, almost perfectly square-shaped skull up to 22 centimetres (8.7 in) in both width and length. [2] The squared shape is caused primarily by a widening of the paired nasal bones along the midline of the snout. The premaxilla bones at the tip of the snout form the entire front edge of the square, while the maxilla bones form the side edges. The maxillae of Spathicephalus are unusual in being thin (no more than 1 centimetre (0.39 in) in thickness) along their entire length. The orbits or eye sockets are large, facing directly upward and positioned close together near the back of the skull. Unlike the roughly circular orbits of most tetrapods, the orbits of Spathicephalus are kidney-shaped because they have fused with another pair of holes called the antorbital fenestrae (in other early tetrapods, the antorbital fenestrae lie in front of the orbits and are separate from them). The skull table behind the orbits is very small in comparison to those of other Carboniferous tetrapods, but like those of other tetrapods, it bears a small hole in its center called a pineal foramen. Small tabular "horns" extend laterally from the back of the skull table, forming the temporal notches. The surface of the skull roof is covered by raised bumps and ridges, a condition paleontologists E. H. Beaumont and T. R. Smithson describe as "pustular ornamentation". [2]
Another distinguishing feature of S. mirus is its dentition; it jaws are lined with hundreds of small, chisel-shaped, closely spaced teeth. These marginal teeth are each about 3 millimetres (0.12 in) in cross-sectional diameter and form an unbroken row along both the upper and lower jaws. While they point directly downward in the upper jaw, the marginal teeth slant slightly inward (mesially) in the lower jaw. Teeth on the vomer bones form a second parallel row on the palate that is not as extensive as the marginal tooth row. At about 1 millimetre (0.10 cm) in diameter, these teeth are smaller than the marginal teeth. The rest of the palate has subtle pustular ornamentation at a finer scale than on the skull roof. This is a unique condition among early tetrapods, many of which have more extensive ornamentation on the palate including bony denticles, additional tooth rows, and palatal tusks. The dentition of Spathicephalus is so unusual that isolated jaw fragments with teeth in them can easily be identified. [2]
Although the species S. pereger is only known by a partial skull roof, paleontologist Donald Baird assigned it to Spathicephalus with confidence because the shapes of its bones closely matched the corresponding bones in S. mirus. One of the few differences that separate S. pereger from S. mirus is the reticulate or "honeycomb" ornamentation on its skull roof, characterized by pits and grooves. The pustular ornamentation seen in S. mirus is in fact rare among early tetrapods (plagiosaurid temnospondyls and the very early tetrapod Ichthyostega are some of the few to have them), while the reticulate ornamentation of S. pereger is more typical. [2]
Spathicephalus is an early member of the group Tetrapoda, which includes all four-limbed vertebrates. According to the most recent studies of early tetrapod phylogeny, it belongs to a clade or evolutionary grouping within Tetrapoda called Baphetoidea but lies outside a subgroup of baphetoids called Baphetidae, which form the "core" of Baphetoidea. Most recent studies of tetrapod evolutionary relationships find baphetoids to be stem group tetrapods, meaning that they branched off from other tetrapods before the most recent common ancestor of living tetrapods (amphibians, reptiles, birds, and mammals) appeared. Paleontologists recognized that Spathicephalus was a close relative of baphetids ever since D. M. S. Watson first described it in 1929. Two main features link Spathicephalus with baphetids: antorbital fenestrae that have fused with the orbits, and a closed palate formed mostly from a pair of broad pterygoid bones. Watson placed Spathicephalus in Loxommatidae, a family he had named twelve years earlier to include what are now called baphetids. Although the name Baphetidae has existed since 1865 and takes precedence over Loxommatidae, paleontologists referred to these tetrapods as loxommids throughout much of the twentieth century. In a 1947 review of early tetrapods, paleontologist Alfred Romer called Spathicephalus "a grotesque type of loxommid," considering its unusual skull to be evidence of a primitive position within the group. [5]
Spathicephalus was not included in a modern phylogenetic analysis of early tetrapod relationships until the 2000s. In 2009, paleontologists Angela Milner, Andrew Milner, and Stig Walsh incorporated the anatomical characteristics of S. mirus into an analysis of baphetoid interrelationships. The analysis placed Spathicephalus just outside Baphetidae as the sister taxon of the group, a result which they used to justify its placement in a distinct family, Spathicephalidae. The analysis also found Eucritta melanolimnetes , an older species from the Viséan stage of Scotland, to be the most basal member of Baphetoidea. Below is a cladogram showing the results of their analysis: [6]
Tetrapoda |
| ||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
The enigmatic Carboniferous tetrapod species Doragnathus woodi may be related to Spathicephalus. In two papers published in 1993 and 1994, Russian paleontologist O. A. Lebedev proposed that Doragnathus was a junior synonym of Spathicephalus. [7] [8] Doragnathus woodi, named by Smithson in 1980 from the Dora Open Cast Mine in Fife, Scotland (the same mine where paleontologist Stanley P. Wood discovered remains of S. mirus), is known only from upper and lower jaw fragments. [9] The jaws themselves closely resemble those of other Carboniferous tetrapods, but the numerous very small, closely packed, pointed, and inward (lingually) curving marginal teeth they hold are unique. Given that Doragnathus and Spathicephalus were the only known Carboniferous tetrapods at the time with marginal teeth that are very small and closely packed, Lebedev argued that the two represent the same taxon. Because the Doragnathus material was smaller than every S. mirus specimen that was known at the time, he also proposed that Doragnathus represented the juvenile form of Spathicephalus and that the curved, pointed teeth of the former transformed into the straight, chizel-shaped teeth of the latter during development. However, this hypothesis was refuted by Beaumont and Smithson, who in 1998 reported a jaw of S. mirus that was just as small as the Doragnathus material but possessed straight, chizel-shaped teeth. Beaumont and Smithson also pointed out that Doragnathus differs from Spathicephalus in possessing an additional tooth row along the parasymphesial plate and coronoids (a series of bones forming the inner parts of the lower jaw of early tetrapods that was lost later in tetrapod evolution). [2]
Another stem tetrapod, Sigournea multidentata from the Early Carboniferous of the United States, may also be related to Spathicephalus. Named in 2006 from a Visean-aged fissure fill deposit in Iowa, Sigournea is slightly older than Spathicephalus. It resembles both Spathicephalus and Doragnathus in having numerous small, closely packed teeth. Sigournea differs from Spathicephalus and resembles Doragnathus in having pointed rather than chizel-shaped marginal teeth and a second row of teeth in the lower jaws, and differs from both taxa in having a hole on the inner surface of the lower jaw called the exomeckelian fenestra. However, given that Sigournea is only known from a single lower jaw, its relationships to other tetrapods remain uncertain. [10]
Milner et al. (2009) raised the possibility that Doragnathus woodi and Sigournea multidentata could both belong to Spathicephalidae, although they did not include the two species in their phylogenetic analysis. [6] An earlier phylogenetic analysis published by Marcello Ruta and John R. Bolt in 2008 included Doragnathus woodi, Sigournea multidentata, and Spathicephalus mirus but did not find all three species to group together. Instead they found good support for Sigournea grouping with Occidens portlocki from the earliest Carboniferous (Tournaisian) of Ireland and Doragnathus grouping with Greererpeton burkemorani from the Viséan of West Virginia in the United States. [11]
The bizarre cranial morphology of Spathicephalus suggests that its feeding habits differed greatly from those of other Carboniferous tetrapods. Most stem tetrapods including baphetids were most likely piscivorous (fish-eating) given their large teeth and deep skulls, which provided attachment points for strong, fast-acting jaw muscles. The small chizel-shaped teeth of Spathicephalus would have been ill-suited for catching fish. Moreover, the flattened shape of its skull means that the depressor mandibulae, muscles that attach to the back of the skull and are responsible for opening the lower jaw, would not have had much room to anchor and therefore would have had poor mechanical advantage. [2]
Although only distantly related, another group of flat-headed aquatic tetrapods called plagiosaurid temnospondyls may have resembled Spathicephalus in their feeding behavior. Plagiosaurids, which lived during the Triassic, are thought to have been bottom-dwelling piscivores that compensated for their weak depressor mandibulae with powerful adductor muscles that would have rapidly closed the lower jaw. They may have rested on the bottom of lakes and rivers with their jaws open, snapping them shut to capture fish. [12] Paleontologist Alfred Romer proposed in 1947 that Spathicephalus was a bottom-dwelling piscivore, but unlike plagiosaurids it would have had weak adductor muscles. [5] In Spathicephalus, plagiosaurids, and many other early tetrapods, a pair of holes at the back of the palate called the subtemporal fossae function as anchors for the adductors. These fossae are large in plagiosaurids, which is why they are thought to have had powerful adductors. Spathicephalus, however, has smaller subtemporal fossae, meaning that it would have had weak adductors and depressor mandibulae. In 1998, paleontologists E. H. Beaumont and T. R. Smithson hypothesized that Spathicephalus fed on aquatic invertebrates through a combination of suction feeding and filter feeding. To feed, it would have rested on a lake or river bottom with its jaws slightly opened and its hyoid apparatus closing off the throat. When a group of small invertebrates approached, it would have lowered its hyoid, opening the throat and causing a rapid decrease in pressure inside the mouth. Water would then flow into the mouth, carrying the prey with it. Even with weak adductors, Spathicephalus could have shut its jaws before prey escaped, and then raised the hyoid again to force water back out while filtering the invertebrates with its rows of small teeth. [2]
Probainognathus meaning “progressive jaw” is an extinct genus of cynodonts that lived around 235 to 221.5 million years ago, during the Late Triassic in what is now Argentina. Together with the genus Bonacynodon from Brazil, Probainognathus forms the family Probainognathidae. Probainognathus was a relatively small, carnivorous or insectivorous cynodont. Like all cynodonts, it was a relative of mammals, and it possessed several mammal-like features. Like some other cynodonts, Probainognathus had a double jaw joint, which not only included the quadrate and articular bones like in more basal synapsids, but also the squamosal and surangular bones. A joint between the dentary and squamosal bones, as seen in modern mammals, was however absent in Probainognathus.
Temnospondyli or temnospondyls is a diverse ancient order of small to giant tetrapods — often considered primitive amphibians — that flourished worldwide during the Carboniferous, Permian and Triassic periods, with fossils being found on every continent. A few species continued into the Jurassic and Early Cretaceous periods, but all had gone extinct by the Late Cretaceous. During about 210 million years of evolutionary history, they adapted to a wide range of habitats, including freshwater, terrestrial, and even coastal marine environments. Their life history is well understood, with fossils known from the larval stage, metamorphosis, and maturity. Most temnospondyls were semiaquatic, although some were almost fully terrestrial, returning to the water only to breed. These temnospondyls were some of the first vertebrates fully adapted to life on land. Although temnospondyls are amphibians, many had characteristics such as scales and armour-like bony plates that distinguish them from the modern soft-bodied lissamphibians.
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. Members of the genus bore a superficial resemblance to modern-day lizards. The genus is known from a single species, Westlothiana lizziae. The type specimen was discovered in the East Kirkton Limestone at the East Kirkton Quarry, West Lothian, Scotland in 1984. This specimen was nicknamed "Lizzie the lizard" by fossil hunter Stan Wood, and this name was quickly adopted by other paleontologists and the press. When the specimen was formally named in 1990, it was given the specific name "lizziae" in homage to this nickname. However, despite its similar body shape, Westlothiana is not considered a true lizard. Westlothiana's anatomy contained a mixture of both "labyrinthodont" and reptilian features, and was originally regarded as the oldest known reptile or amniote. However, updated studies have shown that this identification is not entirely accurate. Instead of being one of the first amniotes, Westlothiana was rather a close relative of Amniota. As a result, most paleontologists since the original description place the genus within the group Reptiliomorpha, among other amniote relatives such as diadectomorphs and seymouriamorphs. Later analyses usually place the genus as the earliest diverging member of Lepospondyli, a collection of unusual tetrapods which may be close to amniotes or lissamphibians, or potentially both at the same time.
Ophiacodon is an extinct genus of synapsid belonging to the family Ophiacodontidae that lived from the Late Carboniferous to the Early Permian in North America and Europe. The genus was named along with its type species O. mirus by paleontologist Othniel Charles Marsh in 1878 and currently includes five other species. As an ophiacodontid, Ophiacodon is one of the most basal synapsids and is close to the evolutionary line leading to mammals.
Secodontosaurus is an extinct genus of "pelycosaur" synapsids that lived from between about 285 to 272 million years ago during the Early Permian. Like the well known Dimetrodon, Secodontosaurus is a carnivorous member of the Eupelycosauria family Sphenacodontidae and has a similar tall dorsal sail. However, its skull is long, low, and narrow, with slender jaws that have teeth that are very similar in size and shape—unlike the shorter, deep skull of Dimetrodon, which has large, prominent canine-like teeth in front and smaller slicing teeth further back in its jaws. Its unusual long, narrow jaws suggest that Secodontosaurus may have been specialized for catching fish or for hunting prey that lived or hid in burrows or crevices. Although no complete skeletons are currently known, Secodontosaurus likely ranged from about 2 to 2.7 metres (7–9 ft) in length, weighing up to 110 kilograms (250 lb).
Crassigyrinus is an extinct genus of carnivorous stem tetrapod from the Early Carboniferous Limestone Coal Group of Scotland and possibly Greer, West Virginia.
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.
Mastodonsaurus is an extinct genus of temnospondyl amphibian from the Middle Triassic of Europe. It belongs to a Triassic group of temnospondyls called Capitosauria, characterized by their large body size and presumably aquatic lifestyles. Mastodonsaurus remains one of the largest amphibians known, and may have exceeded 6 meters in length.
Colosteidae is a family of stegocephalians that lived in the Carboniferous period. They possessed a variety of characteristics from different tetrapod or stem-tetrapod groups, which made them historically difficult to classify. They are now considered to be part of a lineage intermediate between the earliest Devonian terrestrial vertebrates, and the different groups ancestral to all modern tetrapods, such as temnospondyls and reptiliomorphs.
Saharastega is an extinct genus of basal temnospondyl which lived during the Late Permian period, around 251 to 260 million years ago. Remains of Saharastega, discovered by paleontologist Christian Sidor at the Moradi Formation in Niger, were described briefly in 2005 and more comprehensively in 2006. The description is based on a skull lacking the lower jaws.
Loxomma is an extinct genus of Loxommatinae and one of the first Carboniferous tetrapods. They were first described in 1862 and further described in 1870 when two more craniums were found. It is mostly associated with the area of the United Kingdom. They share features with modern reptiles as well as with fish. They had 4 paddle-like limbs that they used to swim in lakes, but they breathed air. Their diet consisted mostly of live fish. They are of the family Baphetidae which are distinguished by their keyhole shaped orbits, while Loxomma themselves are distinguished by the unique texture on their skulls, said to be honeycomb-like.
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.
Boii is an extinct genus of microsaur within the family Tuditanidae. 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. For many years the type and only known species, Boii crassidens, was considered to be a species of Sparodus, until 1966 when Robert Carroll assigned it to its own genus.
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.
Kyrinion is an extinct genus of baphetid tetrapod from the Late Carboniferous of England. It is known from a skull that was found in Tyne and Wear county dating back to the Westphalian stage. Along with the skull is part of the lower jaw, an arch of the atlas bone and a rib possibly belonging to a cervical (neck) vertebra. The type species K. martilli was named from this material in 2003.
Whatcheeria is an extinct genus of early tetrapod from the Mississippian of Iowa. Fossils have been found in 340 million year old fissure fill deposits in the town of Delta. The type species, Whatcheeria deltae was named in 1995. It is classified within the family Whatcheeriidae, along with the closely related Pederpes and possibly Ossinodus.
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.
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.
Occidens is an extinct genus of stem tetrapod from the Early Carboniferous (Tournaisian) Altagoan Formation of Northern Ireland. It is known from a single type species, Occidens portlocki, named in 2004 on the basis of a left lower jaw described by British geologist Joseph Ellison Portlock in 1843.
Gordodon is an extinct genus of non-mammalian synapsid that lived during the Early Permian of what is now Otero County, New Mexico. It was a member of the herbivorous sail-backed family Edaphosauridae and contains only a single species, the type species G. kraineri. Gordodon is unusual among early synapsids for its teeth, which were arranged similarly to those of modern mammals and unlike the simple, uniform lizard-like teeth of other early herbivorous synapsids. Gordodon had large incisor-like teeth at the front, followed by a prominent gap between them and a short row of peg-like teeth at the back. Gordodon was also relatively long-necked for an early synapsid, with elongated and gracile vertebrae in its neck and back. Like other edaphosaurids, Gordodon had a tall sail on its back made from the bony neural spines of its vertebrae. The spines also had bony knobs on them, a common trait of edaphosaurids, but the knobs of Gordodon are also unique for being more slender, thorn-like and randomly arranged along the spines. It is estimated to have been rather small at 1 m in length excluding the tail and 34 kg (75 lb) in weight.