Whatcheeria Temporal range: Early Carboniferous, | |
---|---|
Skull diagram | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Sarcopterygii |
Clade: | Tetrapodomorpha |
Clade: | Elpistostegalia |
Clade: | Stegocephali |
Family: | † Whatcheeriidae |
Genus: | † Whatcheeria Lombard and Bolt, 1995 |
Species | |
|
Whatcheeria is an extinct genus of early tetrapod from the Mississippian (Early Carboniferous) 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. [1] [2] [3]
Whatcheeria is named after What Cheer, Iowa, the hometown of Pat McAdams, the geologist who discovered the first skeletons of the animal. The species is named after Delta, Iowa, the location where the fossils were uncovered. [1]
Whatcheeria possesses a mixture of both primitive and derived traits. It shares with earlier stem tetrapods a series of lateral lines across the skull, rows of teeth on the palate, and small Meckelian foramina across the surface of the lower jaw. It has a cleithrum, a bone in the pectoral girdle that extends from the scapula. The cleithrum once attached to the skull in lobe-finned fish, the ancestors of tetrapods, but detached to allow the neck to move freely. [1]
Whatcheeria grew to up to 2 metres (6.6 ft) long. [2] The skull is deep and the snout is pointed. A hole on the top of the skull behind the eyes called the parietal foramen is relatively large in Whatcheeria. The bones on the skull surface are unusually smooth, unlike the pitted skulls of many other early tetrapods. In front of the eye socket, the prefrontal bone forms a prominent ridge. The prefrontal also projects downward to cover a possible sinus. [1]
The Delta fossil locality was discovered at the abandoned Jasper Hiemstra Quarry and excavated by paleontologists in the late 1980s. [4] The quarry preserves a pair of prehistoric bowl-shaped sinkhole deposits which collapsed into underlying muddy limestone layers of the Waugh and Verdi members of the St. Louis Formation. Most tetrapod fossils are concentrated in a narrow band of fine limy conglomerate deposited above a package of coarser breccia (from the initial collapse) and below the resurgence of muddy limestone. Invertebrate microfossils indicate that these sinkhole deposits are from the Mississippian (Early Carboniferous) subperiod. More precisely, they are probably late Viséan in age (Asbian, stage V3b). This corresponds to a maximum of 340-335 [1] [4] and a minimum of 333-326 million years old [5] (possibly as young as the early Serpukhovian). [2] The geology of the site conforms with an isolated brackish water environment, such as a lagoon, estuary, [1] [4] or mostly freshwater lake with only occasional marine influences. [5]
Over 600 tetrapod fossils have been found at the Delta locality, some of which are partially articulated skulls and skeletons. [1] Whatcheeria is estimated to represent up to 90% of tetrapod fossils from the site, [1] though only 26 specimens can be referred to the genus with certainty. [2] Prior to its 1995 description, Whatcheeria was referred to as a "proto- anthracosaur". [4] Other tetrapods include the colosteid Deltaherpeton [6] and an undescribed species of embolomere. [1] Freshwater fish fossils are common, [4] [1] [5] including remains of rhizodonts, putative "osteolepiforms", "palaeoniscoids", xenacanth sharks, petalodont-like chondrichthyans, [4] [5] gyracanthid acanthodians, [4] and the lungfish Tranodis . [1] Ostracods, snails, myriapods, [5] and plant fossils have also been found at the site. [1]
Whatcheeria was likely primarily aquatic: the poorly-ossified ankle and wrist are ill-suited for locomotion on land, as are the simple, blocky phalanges. This is supported further by the presence of lateral line canals on the skull. Regardless, the limbs are very large and strongly-built, with the shape of the humerus and ulna emphasizing retraction (swinging the forearms back to the torso) over any other direction of movement. Walking would have required strong lateral flexion (sideways bending) of the spine to allow the arms enough freedom of movement. Regardless, Whatcheeria was probably capable of significant lateral flexion due to its unspecialized posterior torso, similar to other early tetrapods. When swimming, Whatcheeria likely used its protruding limbs and paddle-like hands more than its short body or tail. A modern analogue may be the duckbilled platypus (Ornithorhynchus anatinus), a mammal which swims at low speed but high maneuverability via a paddling motion of the forelimbs. In life, Whatcheeria potentially hunted by walking along lakebeds or wading through shallow water, using its relatively flexible neck to augment its ability to capture prey. [2] University of Chicago paleobiologists have likened its estimated lifestyle to modern freshwater predatory reptiles such as crocodilians and the alligator snapping turtle. [7]
The unusually narrow skull of Whatcheeria was strongly reinforced by complex modes of contact between its constituent bones, similar to the Devonian tetrapod Acanthostega . The rear of the skull was replete with interdigitating sutures between the bones of the skull roof and cheek, which would have diluted forces of compression between the sides and top of the rear skull. The snout was supplied with various front-to-back overlapping scarf joints, which would have resisted torsion (twisting) from struggling prey. Whatcheeria emphasizes these traits even further than Acanthostega, combining interdigitation and scarf joints at the front of the palate, the tip of the snout, and throughout the lower jaw. This may correspond to greater strength (and thus more reinforcement) at the front of the jaw when attacking prey, a notion supported by larger anterior fangs in Whatcheeria than other early tetrapods. There are no adaptations for cranial kinesis or suction feeding in Whatcheeria; as a whole, its skull was a stable and strong platform for biting, with an emphasis on the front of the snout for initial prey capture. [3]
Fossils of Whatcheeria represent a range of body sizes and ontogenetic stages, allowing it to decipher growth patterns in early tetrapods. Nine femora (thigh bones) from four size classes have been sampled for histological analyses, cutting a cross section through each bone to determine its developmental history. In larger femora, the cortex (hard outer bone layer) becomes proportionally thinner relative to the smallest femora, where the cortex makes up more than half of the bone's volume. [8]
Size-related variation also shows up in the type of bone deposited in each femur. The smallest femora (size classes one and two, late juveniles to subadults) have a mixture of fibrolamellar bone (a fast-developing composite material combining random bone fibers and concreted osteons) and parallel-fibered bone (fibrous layers woven together at a medium rate along the inner cortex). The largest femora (size classes three and four, adults) lose their fibrolamellar bone and gain lamellar bone (dense, plate-like bone slowly deposited along the circumference of the outer cortex).
The presence of fibrolamellar bone is unique to Whatcheeria among early tetrapods, and is an indicator of fast juvenile development more similar to amniotes than most extinct or living amphibians. [8] This condition suggests previously unexpected variability in the development of early tetrapods, considering there is also some evidence for faster-than-expected growth in Eusthenopteron , a tetrapodomorph fish related to tetrapods. [7] The presence of parallel-fibered bone also indicate that the smallest known femora merely represent late juveniles, and that younger individuals, which likely developed even faster, have not been fossilized at the Delta locality. None of the femora have growth marks, meaning that growth was continuous year-round, and not interrupted by resource scarcity or adverse seasonality. A thin cortex may allow adult Whatcheeria to control their buoyancy more precisely, a useful adaptation for a large aquatic predator. Greererpeton , a colosteid from the same general time period, grew slower and less continuously, and retained a thick cortex well into adulthood. These developmental differences may be a consequence of niche differentiation. [8]
Acanthostega is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period about 365 million years ago, and was anatomically intermediate between lobe-finned fishes and those that were fully capable of coming onto land.
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.
Hynerpeton is an extinct genus of early four-limbed vertebrate that lived in the rivers and ponds of Pennsylvania during the Late Devonian period, around 365 to 363 million years ago. The only known species of Hynerpeton is H. bassetti, named after the describer's grandfather, city planner Edward Bassett. Hynerpeton is known for being the first Devonian four-limbed vertebrate discovered in the United States, as well as possibly being one of the first to have lost internal (fish-like) gills.
Crassigyrinus is an extinct genus of carnivorous stem tetrapod from the Early Carboniferous Limestone Coal Group of Scotland and possibly Greer, West Virginia.
Ventastega is an extinct genus of stem tetrapod that lived during the Upper Fammenian of the Late Devonian, approximately 372.2 to 358.9 million years ago. Only one species is known that belongs in the genus, Ventastega curonica, which was described in 1996 after fossils were discovered in 1933 and mistakenly associated with a fish called Polyplocodus wenjukovi. ‘Curonica’ in the species name refers to Curonia, the Latin name for Kurzeme, a region in western Latvia. Ventastega curonica was discovered in two localities in Latvia, and was the first stem tetrapod described in Latvia along with being only the 4th Devonian tetrapodomorph known at the time of description. Based on the morphology of both cranial and post-cranial elements discovered, Ventastega is more primitive than other Devonian tetrapodomorphs including Acanthostega and Ichthyostega, and helps further understanding of the fish-tetrapod transition.
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.
Tulerpeton is an extinct genus of Devonian four-limbed vertebrate, known from a fossil that was found in the Tula Region of Russia at a site named Andreyevka. This genus and the closely related Acanthostega and Ichthyostega represent the earliest tetrapods.
Cochleosaurus (“spoon lizard”, from the Latin cochlear "spoon" and Greek sauros “lizard”_ were medium-sized edopoid temnospondyls that lived in Euramerica during the Muscovian period. Two species, C. bohemicus and C. florensis, have been identified from the fossil record.
Pederpes is an extinct genus of early Carboniferous tetrapod, dating from 348 to 347.6 Ma in the Tournaisian age. Pederpes contains one species, P. finneyae, 1 m long.
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.
Spathicephalus is an extinct genus of stem tetrapods 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.
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.
Odontocyclops is an extinct genus of Dicynodonts that lived in the Late Permian. Dicynodonts are believed to be the first major assemblage of terrestrial herbivores. Fossils of Odontocyclops have been found in the Karoo Basin of South Africa and the Luangwa Valley of Zambia. The phylogenetic classification of Odontocyclops has been long under debate, but most current research places them as their own genus of Dicynodonts and being very closely related to Rhachiocephalus and Oudenodon.
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.
Deltaherpeton is an extinct genus of colosteid from middle Mississippian deposits of Delta, Iowa, United States. It was first named by John R. Bolt and R. Eric Lombard in 2010 and the type species is Deltaherpeton hiemstrae.
Antlerpeton is an extinct genus of early tetrapod from the Early Carboniferous of Nevada. It is known from a single poorly preserved skeleton from the Diamond Peak Formation of Eureka County. A mix of features in its compound vertebrae suggest that Antlerpeton is a primitive stem tetrapod that has affinities with later, more advanced forms. Its robust pelvis and hind limbs allowed for effective locomotion on land, but the animal was likely still tied to a semiaquatic lifestyle near the coast.
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 amphibian 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.
Innovations conventionally associated with terrestrially first appeared in aquatic elpistostegalians such as Panderichthys rhombolepis, Elpistostege watsoni, and Tiktaalik roseae. Phylogenetic analyses distribute the features that developed along the tetrapod stem and display a stepwise process of character acquisition, rather than abrupt. The complete transition occurred over a period of 30 million years beginning with the tetrapodomorph diversification in the Middle Devonian.
The Intertemporal bone is a paired cranial bone present in certain sarcopterygians and extinct amphibian-grade tetrapods. It lies in the rear part of the skull, behind the eyes.