Aistopoda (Greek for " avingnot-visible feet") is an order of highly specialised snake-like stegocephalians known from the

Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...

and

Early Permian 01 or 01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), ''01'' (Richard Müller album), 2001 * 01 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...

of Europe and North America, ranging from tiny forms only , to nearly in length. They first appear in the fossil record in the Mississippian period and continue through to the

. The

skull The skull, or cranium, is typically a bony enclosure around the brain of a vertebrate. In some fish, and amphibians, the skull is of cartilage. The skull is at the head end of the vertebrate. In the human, the skull comprises two prominent ...

is small but very specialised, with large

orbits In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...

, and large

fenestra A fenestra (fenestration; : fenestrae or fenestrations) is any small opening or pore, commonly used as a term in the biology, biological sciences. It is the Latin word for "window", and is used in various fields to describe a pore in an anatomy, ...

e. The primitive form ''

Ophiderpeton ''Ophiderpeton'' (from , 'snake' and 'creeper') is an extinct genus of aistopod tetrapodomorphs from the early Carboniferous to the early Permian. Remains of this genus are widespread and were found in Ohio, United States, Ireland, and the Cz ...

'' has a pattern of

dermal bone A dermal bone or investing bone or membrane bone is a bony structure derived from intramembranous ossification forming components of the vertebrate skeleton, including much of the skull, jaws, gill covers, shoulder girdle, fin rays ( lepidotrich ...

s in the skull similar in respects to the

temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...

s. But in the advanced genus '' Phlegethontia'' the skull is very light and open, reduced to a series of struts supporting the braincase against the lower jaw, just as in

snakes Snakes are elongated Limbless vertebrate, limbless reptiles of the suborder Serpentes (). Cladistically Squamata, squamates, snakes are ectothermic, amniote vertebrates covered in overlapping Scale (zoology), scales much like other members of ...

, and it is possible that the aistopods filled the same

ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of Resource (biology), resources an ...

s in the Paleozoic that snakes do today. Pseudophlegethontia turnbullorum

They had an extremely elongated body, with up to 230

vertebra Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spina ...

e. The vertebrae were holospondylous, having only a single ossification per segment. They lacked intercentra, even in the tail, and had are no free

haemal arch A haemal arch, also known as a chevron, is a bony arch on the ventral side of a tail vertebra of a vertebrate. The canal formed by the space between the arch and the vertebral body is the haemal canal. A spinous ventral process emerging from the ha ...

es. The

neural arch Each vertebra (: vertebrae) is an irregular bone with a complex structure composed of bone and some hyaline cartilage, that make up the vertebral column or spine, of vertebrates. The proportions of the vertebrae differ according to their spinal ...

was low and fused to the centrum. All of these features are very similar to those of the

Nectridea Nectridea is an extinct order of lepospondyl tetrapods from the Carboniferous and Permian periods, including animals such as '' Diplocaulus''. In appearance, they would have resembled modern newts or aquatic salamanders, although they are not cl ...

, both representing the typical lepospondylous condition. A recent paper described the internal organization of the aistopod head, finding that aïstopods retained many fish-like features of the skull and brain, including a persistent extension of the

notochord The notochord is an elastic, rod-like structure found in chordates. In vertebrates the notochord is an embryonic structure that disintegrates, as the vertebrae develop, to become the nucleus pulposus in the intervertebral discs of the verteb ...

into the head and an open canal between the

pituitary gland The pituitary gland or hypophysis is an endocrine gland in vertebrates. In humans, the pituitary gland is located at the base of the human brain, brain, protruding off the bottom of the hypothalamus. The pituitary gland and the hypothalamus contr ...

and the mouth. Because these features are lost early in tetrapod evolution, this may be evidence that aistopods diverged from other tetrapods soon after the origin of digits. The ribs were slender, either single or double-headed, with the head shaped like a letter "K". There is no trace of limbs or even limb girdles in any known fossil, and the tail was short and primitive.

Relationships

Evolutionary relationships with other early

tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...

s remain controversial, as even the earliest aistopod, the

Viséan The Visean, Viséan or Visian is an age in the ICS geologic timescale or a stage in the stratigraphic column. It is the second stage of the Mississippian, the lower subsystem of the Carboniferous. The Visean lasted from to Ma. It follows ...

species '' Lethiscus stocki'', was already highly specialised, although its skull was primitive and extremely fishlike.Fossil holds new insights into how fish evolved onto land
/ref> Aïstopods have been variously grouped with other

lepospondyls Lepospondyli is a diverse clade of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco ('' Diplocaulus minimus''), lepospondyls lived from the Visean stage of the Early Carboniferous to the Earl ...

, or placed at or prior to the batrachomorph-

reptiliomorph Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians (lissamphibians). It was defi ...

divide. However, a cladistic analysis by Pardo ''et al.'' (2017) recovered Aistopoda at the base of Tetrapoda.Jason D. Pardo, Matt Szostakiwskyj, Per E. Ahlberg & Jason S. Anderson (2017) Hidden morphological diversity among early tetrapods. Nature (advance online publication) doi:10.1038/nature22966> The group was quite diverse during the

Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...

, with a few forms continuing through to the

Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...

. Below is a

cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...

from Anderson ''et al.'' (2003) showing the

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

relationships of aistopods:

References

* Benton, M. J. (2000), ''Vertebrate Paleontology'', 2nd ed. Blackwell Science Ltd * Carroll, RL (1988), ''Vertebrate Paleontology and Evolution'', WH Freeman & Co. pp. 176–7 * Reisz, Rober
Biology 356 - Major Features of Vertebrate Evolution
(online) * von Zittel, K.A (1932), ''Textbook of Paleontology'', CR Eastman (transl. and ed), 2nd edition, vol.2, p. 221-2, Macmillan & Co.

External links

{{Authority control Tetrapodomorph orders Aistopoda