Acanthostega (meaning "spiny roof") is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period (Famennian age) about 365 million years ago, and was anatomically intermediate between lobe-finned fishes and those that were fully capable of coming onto land.[1][2]
The 60cm (24in)Acanthostegahad eight digits on each hand (the number of digits on the feet is unclear) linked by webbing, it lacked wrists, and was generally poorly adapted for walking on land. It also had a remarkably fish-like shoulder and forelimb.[3] The front limbs of Acanthostega could not bend forward at the elbow, and therefore could not be brought into a weight bearing position, appearing to be more suitable for paddling or for holding on to aquatic plants. Acanthostega is the earliest stem-tetrapod to show the shift in locomotory dominance from the pectoral girdle to the pelvic girdle.[4]
There are many morphological changes that allowed the pelvic girdle of Acanthostega to become a weight-bearing structure. In more ancestral states the two sides of the girdle were not attached. In Acanthostega there is contact between the two sides and fusion of the girdle with the sacral rib of the vertebral column. These fusions would have made the pelvic region more powerful and equipped to counter the force of gravity when not supported by the buoyancy of an aquatic environment.[4] It had internal gills that were covered like those of fish. It also had lungs, but its ribs were too short to support its chest cavity out of water.[2]
Classification
An illustration of the Devonian speciation of lobe-finned fish
Acanthostega is seen as part of widespread evolutionary radiation in the late Devonian period, starting with purely aquatic finned tetrapodomorphs, with their successors showing increased air-breathing capability and related adaptions to the jaws and gills, as well as more muscular neck allowing freer movement of the head than fish have, and use of the fins to raise the body of the fish.[2] These features are displayed by the earlier Tiktaalik, which like Ichthyostega showed signs of greater abilities to move around on land, but is thought to have been primarily aquatic.[2]
It has been inferred that Acanthostega probably lived in shallow, weed-choked swamps, its legs apparently being adapted for these specific ecosystems. Apart from the presence of limbs, it was not adapted in any way for walking on land. Jennifer A. Clack interprets this as showing that Acanthostega was primarily an aquatic animal descended from fish that never left the sea, and that the specializations of the tetrapod lineage were exaptations: features which would later be useful for terrestrial life, even if they originated for a different purpose. At that period, deciduous plants were flourishing and annually shedding leaves into the water, attracting small prey into warm oxygen-poor shallows that were difficult for larger fish to swim in; Clack remarks on how the lower jaw of Acanthostega shows a change from those of fish that have two rows of teeth, with a large number of small teeth in the outer row, and two large fangs and some smaller teeth in the inner row. This difference likely corresponds to a shift in stem-tetrapods from feeding exclusively in the water to feeding with the head above water or on land.[2]
Acanthostega AMNH model - head detail
Research based on analysis of the suturemorphology in the skull of Acanthostega indicates that the species was able to bite prey at or near the water's edge. Markey and Marshall compared the skull with the skulls of fish, which use suction feeding as the primary method of prey capture, and creatures known to have used the direct biting on prey typical of terrestrial animals. Their results indicate that Acanthostega was adapted for what they call terrestrial-style feeding, strongly supporting the hypothesis that the terrestrial mode of feeding first emerged in aquatic animals. If correct, this shows an animal specialized for hunting and living in shallow waters in the line between land and water.[5]
Lifestyle
Model reconstruction at the State Museum of Natural History in Stuttgart, Germany
While normally considered more basal than Ichthyostega, it is possible that Acanthostega was actually more derived. Since Acanthostega resembles juvenile Ichthyostega and shows a lot less differences from juveniles to adults than the latter, it has been suggested that Acanthostega might be descended from a neotenic lineage. Although it appears to have spent its whole life in water, its humerus also exhibits traits that resemble those of later, fully terrestrial stem-tetrapods (the humerus in Ichthyostega being somewhat derived from, and homologous with the pectoral and pelvic fin bones of earlier fishes). This could indicate that vertebrates evolved terrestrial traits earlier than previously assumed, and numerous times independently from another.[6] Muscle scars on the forelimbs of Acanthostega were similar to those of crown-tetrapods, suggesting that it evolved from an ancestor that had more terrestrial adaptations than itself.[7]
Development
A histological study of Acanthostega humeri, assisted by synchotron scans, indicates that the animal matured slowly. Some individuals reached sexual maturity (based on a fully ossified humerus) at more than six years of age, and adult fossils are much rarer than juveniles. Late ossification of the humerus supports a fully aquatic lifestyle for Acanthostega. There is barely any correlation between humerus size and maturity, suggesting that there was significant size variation among individuals of the same age. This may be due to competitive pressures, differing adaptive strategies, or even sexual dimorphism. However, the small sample size prevents recognition of a bimodal distribution which could confirm the latter hypothesis.[8]
Clack, J. A. (2009). "The fin to limb transition: new data, interpretations, and hypotheses from paleontology and developmental biology". Annual Review of Earth and Planetary Sciences. 37 (1): 163–179. Bibcode:2009AREPS..37..163C. doi:10.1146/annurev.earth.36.031207.124146.
Laurin M. 2010. How Vertebrates Left the Water. Berkeley: University of California Press.
Steyer J-Sb. 2012. Earth Before the Dinosaurs. Bloomington: Indiana University Press.
A tetrapod is any four-limbed vertebrate animal of the superclass Tetrapoda. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids and synapsids. Some tetrapods such as snakes, legless lizards, and caecilians had evolved to become limbless via mutations of the Hox gene, although some do still have a pair of vestigial spurs that are remnants of the hindlimbs.
Sarcopterygii — sometimes considered synonymous with Crossopterygii — is a taxon of the bony fish known as the lobe-finned fish or sarcopterygians, characterised by prominent muscular limb buds (lobes) within the fins, which are supported by articulated appendicular skeletons. This is in contrast to the other clade of bony fish, the Actinopterygii, which have only skin-covered bony spines (lepidotrichia) supporting the fins.
Ichthyostega is an extinct genus of limbed tetrapodomorphs from the Late Devonian of what is now Greenland. It was among the earliest four-limbed vertebrates ever in the fossil record and was one of the first with weight-bearing adaptations for terrestrial locomotion. Ichthyostega possessed lungs and limbs that helped it navigate through shallow water in swamps. Although Ichthyostega is often labelled a 'tetrapod' because of its limbs and fingers, it evolved long before true crown group tetrapods and could more accurately be referred to as a stegocephalian or stem tetrapod. Likewise, while undoubtedly of amphibian build and habit, it is not a true member of the group in the narrow sense, as the first modern amphibians appeared in the Triassic Period. Until finds of other early stegocephalians and closely related fishes in the late 20th century, Ichthyostega stood alone as a transitional fossil between fish and tetrapods, combining fish and tetrapod features. Newer research has shown that it had an unusual anatomy, functioning more akin to a seal than a salamander, as previously assumed.
Jennifer Alice Clack, was an English palaeontologist and evolutionary biologist. She specialised in the early evolution of tetrapods, specifically studying the "fish to tetrapod" transition: the origin, evolutionary development and radiation of early tetrapods and their relatives among the lobe-finned fishes. She is best known for her book Gaining Ground: the Origin and Early Evolution of Tetrapods, published in 2002 and written with the layperson in mind.
"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.
Panderichthys is a genus of extinct sarcopterygian from the late Devonian period, about 380 Mya. Panderichthys, which was recovered from Frasnian deposits in Latvia, is represented by two species. P. stolbovi is known only from some snout fragments and an incomplete lower jaw. P. rhombolepis is known from several more complete specimens. Although it probably belongs to a sister group of the earliest tetrapods, Panderichthys exhibits a range of features transitional between tristichopterid lobe-fin fishes and early tetrapods. It is named after the German-Baltic paleontologist Christian Heinrich Pander. Possible tetrapod tracks dating back to before the appearance of Panderichthys in the fossil record were reported in 2010, which suggests that Panderichthys is not a direct ancestor of tetrapods, but nonetheless shows the traits that evolved during the fish-tetrapod evolution
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.
Tiktaalik is a monospecific genus of extinct sarcopterygian from the Late Devonian Period, about 375 Mya, having many features akin to those of tetrapods. Tiktaalik is estimated to have had a total length of 1.25–2.75 metres (4.1–9.0 ft) based on various specimens.
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.
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.
A limb is a jointed, muscled appendage of a tetrapod vertebrate animal used for weight-bearing, terrestrial locomotion and physical interaction with other objects. The distalmost portion of a limb is known as its extremity. The limbs' bony endoskeleton, known as the appendicular skeleton, is homologous among all tetrapods, who use their limbs for walking, running and jumping, swimming, climbing, grasping, touching and striking.
Elpistostegalia or Panderichthyida is an order of prehistoric lobe-finned fishes which lived during the Middle Devonian to Late Devonian period. They represent the advanced tetrapodomorph stock, the fishes more closely related to tetrapods than the osteolepiform fishes. The earliest elpistostegalians, combining fishlike and tetrapod-like characters, are sometimes called fishapods, a phrase coined for the advanced elpistostegalian Tiktaalik. Through a strict cladistic view, the order includes the terrestrial tetrapods.
Polydactyly in stem-tetrapods should here be understood as having more than five digits to the finger or foot, a condition that was the natural state of affairs in the earliest stegocephalians during the evolution of terrestriality. The polydactyly in these largely aquatic animals is not to be confused with polydactyly in the medical sense, i.e. it was not an anomaly in the sense it was not a congenital condition of having more than the typical number of digits for a given taxon. Rather, it appears to be a result of the early evolution from a limb with a fin rather than digits.
Ichthyostegalia is an order of extinct amphibians, representing the earliest landliving vertebrates. The group is thus an evolutionary grade rather than a clade. While the group are recognized as having feet rather than fins, most, if not all, had internal gills in adulthood and lived primarily as shallow water fish and spent minimal time on land.
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.
Tinirau is an extinct genus of sarcopterygian fish from the Middle Devonian of Nevada. Although it spent its entire life in the ocean, Tinirau is a stem tetrapod close to the ancestry of land-living vertebrates in the crown group Tetrapoda. Relative to more well-known stem tetrapods, Tinirau is more closely related to Tetrapoda than is Eusthenopteron, but farther from Tetrapoda than is Panderichthys. The type and only species of Tinirau is T. clackae, named in 2012.
The evolution of tetrapods began about 400 million years ago in the Devonian Period with the earliest tetrapods evolved from lobe-finned fishes. Tetrapods are categorized as animals in the biological superclass Tetrapoda, which includes all living and extinct amphibians, reptiles, birds, and mammals. While most species today are terrestrial, little evidence supports the idea that any of the earliest tetrapods could move about on land, as their limbs could not have held their midsections off the ground and the known trackways do not indicate they dragged their bellies around. Presumably, the tracks were made by animals walking along the bottoms of shallow bodies of water. The specific aquatic ancestors of the tetrapods, and the process by which land colonization occurred, remain unclear. They are areas of active research and debate among palaeontologists at present.
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 vertebrate land invasion refers to the transition of vertebrate animals from being aquatic/semiaquatic to predominantly terrestrial during the Late Devonian period. This transition allowed some vertebrates to escape competitive pressure from other aquatic animals and explore niches on land, which eventually established the vertebrates as the dominant terrestrial phylum. Fossils from this period have allowed scientists to identify some of the species that existed during this transition, such as Tiktaalik and Acanthostega. Many of these species were also the first to develop adaptations suited to terrestrial over aquatic life, such as neck mobility, more robust lungs and hindlimb locomotion.
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