Skull roof

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The full complement of bones of the tetrapod skull roof, as seen in the temnospondyl Xenotosuchus Xenotosuchus skull roof.svg
The full complement of bones of the tetrapod skull roof, as seen in the temnospondyl Xenotosuchus

The skull roof or the roofing bones of the skull are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium.

Contents

In comparative anatomy, the term is applied to the whole dermatocranium. [1] In general anatomy, the roofing bones may refer specifically to the bones that form above and alongside the brain and neurocranium (i.e., excluding the marginal upper jaw bones such as the maxilla and premaxilla). [2] In human anatomy, the skull roof often refers specifically to the skullcap.

Origin

Dermal armour in Dunkleosteus, a placoderm. Dunkleosteus skull QM email.jpg
Dermal armour in Dunkleosteus, a placoderm.

Early armoured fish (such as jawless ostracoderms and jawed placoderms) did not have a skull in the common understanding of the word, but instead had a cartilaginous endocranium that was partially open from above. The loose cartilage was topped by dermal bones forming armour. The dermal bones gradually evolved into a fixed unit overlaying the endocranium like a heavy "lid", protecting the animal's head and brain from above. A more or less full shield of fused dermal bones was common in early bony fishes of the Devonian, and particularly well developed in shallow water species. [3]

Cartilaginous fish, such as sharks, have a skeleton which is entirely formed from cartilage. They lack a continuous dermal armour and thus have no proper skull roof.

Bony fishes

Skull of Platycephalichthys, a sarcopterygian. Most of the roofing over the cheek region is formed by the operculum. Platycephalichthys.jpg
Skull of Platycephalichthys , a sarcopterygian. Most of the roofing over the cheek region is formed by the operculum.

In early sarcopterygians ("lobe-finned fish"), the skull roof was composed of numerous bony plates, particularly around the nostrils and behind each eye. The skull proper was joined by the bones of the operculum. The skull itself was composed rather loosely, with a joint between the bones covering the brain and the snout.

The skull roof in lungfish is composed of a number of bony plates that are not readily compared to those found in early amphibians. [4] In most ray-finned fishes, the skull is often reduced to a series of loose elements, and a skull roof as such is not found. [3]

Early tetrapods

The skull roof in Cheliderpeton, a temnospondyl amphibian Cheliderpeton skull2.jpg
The skull roof in Cheliderpeton , a temnospondyl amphibian

As tetrapodomorph fish trended towards greater adaptations for life on land, the skull became more tightly integrated. At the same time, the number of bones were reduced, the skull bones separated from the shoulder girdle, and the operculum disappeared. [1] The earliest limbed tetrapods ("amphibians" in the broad sense) solidified a pattern of plates which formed the basis for that seen in all land-living vertebrates. These early tetrapods (including temnospondyls, embolomeres, and various minor groups) have historically been termed "labyrinthodonts" ("maze teeth") or "stegocephalians" ("roof heads"). Not including marginal or cheek bones (such as the premaxillae, maxillae, jugals, quadratojugals, and squamosals), the skull roof bones established by these early tetrapods include the following:

The skull roof itself formed a continuous cover over the whole of the head, leaving only openings for nostrils (nares), eyes (orbits), and a small parietal eye (also known as a pineal foramen) between the parietal bones. This type of skull was inherited by the first amniotes (fully terrestrial tetrapods), which evolved in the Carboniferous. This type of skull roof without any above openings behind the eyes is called anapsid . Today, the only reptiles with anapsid skulls are turtles, though this is likely a case of secondary loss of the temporal fenestrae. [5]

In modern amphibians such as frogs and salamanders, the skull roof is further reduced and has large openings. Only in caecilians can a full covering skull roof be found, an adaption for burrowing. [6]

The debate over skull roof homology

One of the most persistent debates in 20th-century paleontology was how to homologize the skull roof of fish with that of tetrapods. [7] In practically all tetrapods, the midline of the skull roof comprises at least three pairs of plate-like bones. From front-to-back, these bones are the nasals (top of the snout), frontals (between the eyes), and parietals (behind the frontals). Early tetrapods also possessed postparietals, additional paired or singular bones at the rim of the skull, behind the parietals. [8] Lobe-finned fish and early ray-finned fish usually lack paired bones at the top of the snout, instead presenting a mosaic of smaller plates. Nevertheless, two pairs of bones do consistently occur further back. [9] The front pair are positioned between the eyes and surround a pineal foramen, when it is present. The latter pair are elongated and abut the extrascapular bones, which lie behind the skull. The "traditional" or "orthodox" hypothesis considers these two pairs to be equivalent to the frontal and parietal bones, respectively. This was mainly justified by their position in regards to the eyes and brain, in accordance with mammal anatomy. [10] [11]

An alternative interpretation was proposed by T.S. Westoll (1938, 1943) [12] [13] and A.S. Romer (1941). [14] Their interpretation noted that the tetrapods with a pineal foramen almost always have the hole surrounded by the parietal bones. This would indicate that the "frontal" bones of fish are actually parietal bones. By extension, the "parietals" of fish are actually postparietals, while the tetrapod nasal and frontal bones develop from fused snout ossicles. According to this hypothesis, the eyes shift forwards, the snout expands, and the postorbital region (behind the eyes) contracts at the origin of tetrapods. Newly discovered "transitional" fossils such as Ichthyostega , Elpistostege , and Panderichthys were used as further evidence in support of their interpretation. [12] [13]

The Westoll/Romer hypothesis was initially controversial, with the strongest critiques coming from Scandinavian paleoichthyologists. [15] [16] [10] [11] Proponents of the orthodox hypothesis argued that the "parietals" of fish are too strongly connected to the underlying brain anatomy to justify the W/R scenario, which posits that the "parietals" (= postparietals) diminish into oblivion over the course of evolution. Another concern was that the W/R hypothesis necessitates the complete loss of the extrascapulars in tetrapods. According to the orthodox hypothesis, the tetrapod postparietals develop from extrascapulars which have acquired a stronger connection to the skull. [15] [16] [10] [11] Despite the criticism, the W/R scenario is now accepted by most paleontologists studying the fish-tetrapod transition. [17] [18] [8] [9] [7] Developmental studies show support for the novel evolution of the frontal bones in tetrapods and their lack of homology with the "frontals" of fish. [19] Further tetrapodomorph discoveries, such as Acanthostega and Tiktaalik , [20] provide more concrete evidence for the anatomical shifts hypothesized by Westoll and Romer.

Diapsids and synapsids

Tuatara skull, showing the double openings behind the eye. SphenodonSkull.jpg
Tuatara skull, showing the double openings behind the eye.

In two groups of early amniotes, the skull roof evolved temporal fenestrae to allow for greater movement of the jaw muscles. The two groups evolved the openings independently:

Related Research Articles

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The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone, however two parts are more prominent: the cranium and the mandible. In humans, these two parts are the neurocranium (braincase) and the viscerocranium that includes the mandible as its largest bone. The skull forms the anterior-most portion of the skeleton and is a product of cephalisation—housing the brain, and several sensory structures such as the eyes, ears, nose, and mouth. In humans, these sensory structures are part of the facial skeleton.

<span class="mw-page-title-main">Occipital bone</span> Saucer-shaped membrane bone situated at the back and lower part of the cranium

The occipital bone is a cranial dermal bone and the main bone of the occiput. It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobes of the cerebrum. At the base of the skull in the occipital bone, there is a large oval opening called the foramen magnum, which allows the passage of the spinal cord.

<span class="mw-page-title-main">Frontal bone</span> Bone in the human skull

The frontal bone is a bone in the human skull. The bone consists of two portions. These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, part of the bony orbital cavity holding the eye, and part of the bony part of the nose respectively. The name comes from the Latin word frons.

<span class="mw-page-title-main">Parietal bone</span> Bone in the human skull which, when joined together, forms the sides and roof of the cranium

The parietal bones are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is named from the Latin paries (-ietis), wall.

<span class="mw-page-title-main">Nasal bone</span> Two bones in the bridge of the human nose

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<span class="mw-page-title-main">Fish anatomy</span> Study of the form or morphology of fishes

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<span class="mw-page-title-main">Adelospondyli</span> Extinct order of amphibians

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.

<span class="mw-page-title-main">Parietal eye</span> Part of the epithalamus

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<span class="mw-page-title-main">Branchial arch</span> Bony "loops" present in fish, which support the gills

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<span class="mw-page-title-main">Endocranium</span> The lower and inner parts of the skull

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<span class="mw-page-title-main">Postparietal</span> Fish skull bones

Postparietals are cranial bones present in fish and many tetrapods. Although initially a pair of bones, many lineages possess postparietals which were fused into a single bone. The postparietals were dermal bones situated along the midline of the skull, behind the parietal bones. They formed part of the rear edge of the skull roof, and the lateral edge of each postparietal often contacts the tabular and supratemporal bones. In fish, the postparietals are elongated, typically the largest components of the skull roof. Tetrapods possessed shorter postparietals, which were reduced further and shifted towards the braincase in amniotes. At several points in synapsid evolution, the postparietals fused to each other and the tabulars during embryological development. This fusion produces the interparietal bone, which is inherited by mammals. Postparietals are common in extinct amphibians and early reptiles. However, most living amphibians and living reptiles lack postparietal bones, with a few exceptions.

The supratemporal bone is a paired cranial bone present in many tetrapods and tetrapodomorph fish. It is part of the temporal region, usually lying medial (inwards) relative to the squamosal and lateral (outwards) relative to the parietal and/or postparietal. It may also contact the postorbital or intertemporal, or tabular, when those bones are present.

References

  1. 1 2 Romer, A.S. & T.S. Parsons. 1977. The Vertebrate Body. 5th ed. Saunders, Philadelphia. (6th ed. 1985)
  2. ↑ Kent, George C.; Carr, Robert K. (2001). Comparative Anatomy of the Vertebrates (9th ed.). New York, NY: McGraw-Hill. ISBN   0-07-303869-5.
  3. 1 2 Miller, George C. Kent, Larry (1997). Comparative anatomy of the vertebrates (8th ed.). Dubuque, IA: Wm. C. Brown. ISBN   0697243788.{{cite book}}: CS1 maint: multiple names: authors list (link)
  4. ↑ Carroll, R.L. (1988): Vertebrate Paleontology and Evolution, W.H. Freeman & Co, New York NY, p.148
  5. ↑ deBraga, M. and Rieppel, O. (1997). "Reptile phylogeny and the interrelationships of turtles." Zoological Journal of the Linnean Society, 120: 281-354.
  6. ↑ Trueb, William E. Duellman, Linda Trueb; illustrated by Linda (1994). Biology of amphibians (Johns Hopkins pbk. ed.). Baltimore: Johns Hopkins University Press. ISBN   0-8018-4780-X.{{cite book}}: CS1 maint: multiple names: authors list (link)
  7. 1 2 Schultze, Hans-Peter (2008). "Nomenclature and homologization of cranial bones in actinopterygians". In Arratia, Gloria; Schultze, Hans-Peter; Wilson, Mark V.H. (eds.). Mesozoic Fishes 4: Homology and Phylogeny. München: Verlag Dr. Friedrich Pfeil. pp. 23–48. ISBN   978-3-89937-080-5.
  8. 1 2 Panchen, A. L.; Smithson, T. R. (1987). "Character Diagnosis, Fossils and the Origin of Tetrapods". Biological Reviews. 62 (4): 341–436. doi:10.1111/j.1469-185X.1987.tb01635.x. ISSN   1469-185X. S2CID   83672017.
  9. 1 2 Ahlberg, Per Erik (1991-11-01). "A re-examination of sarcopterygian interrelationships, with special reference to the Porolepiformes". Zoological Journal of the Linnean Society. 103 (3): 241–287. doi:10.1111/j.1096-3642.1991.tb00905.x. ISSN   0024-4082.
  10. 1 2 3 Jarvik, Erik (15 April 1996). "The Devonian tetrapod Ichthyostega" (PDF). Fossils and Strata. 40: 1–206. doi:10.18261/8200376605-1996-01. ISBN   8200376605.
  11. 1 2 3 Borgen, Ulf J.; Nakrem, Hans A. (2016), Morphology, phylogeny and taxonomy of osteolepiform fish, Fossils and Strata, vol. 61, Wiley-Blackwell, pp. 1–481, doi:10.1002/9781119286448.ch1, ISBN   978-1-119-28643-1 , retrieved 2023-09-24
  12. 1 2 Westoll, T. Stanley (1938). "Ancestry of the Tetrapods". Nature. 141 (3559): 127–128. Bibcode:1938Natur.141..127W. doi:10.1038/141127a0. ISSN   1476-4687. S2CID   4086668.
  13. 1 2 Westoll, T. S. (1943). "The Origin of the Tetrapods". Biological Reviews. 18 (2): 78–98. doi:10.1111/j.1469-185X.1943.tb00289.x. ISSN   1464-7931. S2CID   85648376.
  14. ↑ Romer, Alfred Sherwood (1941). "Notes on the crossopterygian hyomandibular and braincase". Journal of Morphology. 69 (1): 141–160. doi:10.1002/jmor.1050690106. ISSN   0362-2525. S2CID   85150870.
  15. 1 2 Jarvik, E. (1967). "The homology of frontal and parietal bones in fishes and tetrapods". Coll. Internatn. C.N.R.S. 163: 181–213.
  16. 1 2 Borgen, Ulf J. (1983). "Homologizations of skull roofing bones between tetrapods and osteolepiform fishes" (PDF). Paleontology. 26 (4): 735–753.
  17. ↑ Parrington, F. R. (1967-10-01). "The identification of the dermal bones of the head". Zoological Journal of the Linnean Society. 47 (311): 231–239. doi:10.1111/j.1096-3642.1967.tb01406.x. ISSN   0024-4082.
  18. ↑ Schultze, H.P.; Arsenault, M. (1985). "The Panderichthyid fish Elpistostege: a close relative of tetrapods?" (PDF). Paleontology. 28 (2): 293–309.
  19. ↑ Teng, Camilla S; Cavin, Lionel; Maxson, Robert E Jnr; Sánchez-Villagra, Marcelo R; Crump, J Gage (2019-12-23). Tautz, Diethard (ed.). "Resolving homology in the face of shifting germ layer origins: Lessons from a major skull vault boundary". eLife. 8: e52814. doi: 10.7554/eLife.52814 . ISSN   2050-084X. PMC   6927740 . PMID   31869306.
  20. ↑ Edward B. Daeschler, Neil H. Shubin and Farish A. Jenkins Jr. (6 April 2006). "A Devonian tetrapod-like fish and the evolution of the tetrapod body plan" (PDF). Nature . 440 (7085): 757–763. Bibcode:2006Natur.440..757D. doi: 10.1038/nature04639 . PMID   16598249.
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