{{short description|Roofing bones of the skull}} [[File:Xenotosuchus skull roof.svg|thumb|right|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, including land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium.
In comparative anatomy, the term is applied to the whole dermatocranium.<ref name="Romer, A 1977">Romer, A.S. & T.S. Parsons. 1977. ''The Vertebrate Body.'' 5th ed. Saunders, Philadelphia. (6th ed. 1985)</ref> 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).<ref name=kentcarr>{{cite book |last1=Kent |first1=George C.|last2=Carr |first2=Robert K.|year=2001|title=Comparative Anatomy of the Vertebrates|edition=9th |location=New York, NY |publisher=McGraw-Hill |isbn= 0-07-303869-5}}</ref> In human anatomy, the skull roof often refers specifically to the skullcap.
==Origin== thumb|right|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.<ref name=Kent&Miller>{{cite book|last=Miller|first=George C. Kent, Larry|title=Comparative anatomy of the vertebrates|year=1997|publisher=Wm. C. Brown|location=Dubuque, IA|isbn=0-697-24378-8|edition=8th}}</ref>
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== [[File:Platycephalichthys.jpg|thumb|left|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.<ref>Carroll, R.L. (1988): Vertebrate Paleontology and Evolution,'' W.H. Freeman & Co'', New York NY, p.148</ref> 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.<ref name=Kent&Miller/>
==Early tetrapods== [[File:Cheliderpeton skull2.jpg|thumb|right|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.<ref name="Romer, A 1977" /> 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:
* Nasals (midline) * Lacrimals * Prefrontals * Frontals (midline) * Postfrontals * Postorbitals * Parietals (midline) * Intertemporals * Supratemporals * Postparietals (midline) * Tabulars
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.<ref name="debraga&rieppel1997">deBraga, M. and Rieppel, O. (1997). "Reptile phylogeny and the interrelationships of turtles." ''Zoological Journal of the Linnean Society'', '''120''': 281-354.</ref>
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.<ref name="Duellman&Trueb">{{cite book |last=Duellman |first=William E. |last2=Trueb |first2=Linda |author-link2=Linda Trueb |title=Biology of amphibians |publisher=Johns Hopkins University Press |year=1994 |isbn=0-8018-4780-X |edition=Johns Hopkins pbk. |location=Baltimore}}</ref>
=== 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.<ref name=":1">{{Cite book |last=Schultze |first=Hans-Peter |title=Mesozoic Fishes 4: Homology and Phylogeny |publisher=Verlag Dr. Friedrich Pfeil |year=2008 |isbn=978-3-89937-080-5 |editor-last=Arratia |editor-first=Gloria |location=München |pages=23–48 |chapter=Nomenclature and homologization of cranial bones in actinopterygians |editor-last2=Schultze |editor-first2=Hans-Peter |editor-last3=Wilson |editor-first3=Mark V.H. |chapter-url=https://www.researchgate.net/publication/259334578}}</ref> 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.<ref name=":2">{{Cite journal |last1=Panchen |first1=A. L. |last2=Smithson |first2=T. R. |date=1987 |title=Character Diagnosis, Fossils and the Origin of Tetrapods |journal=Biological Reviews |language=en |volume=62 |issue=4 |pages=341–436 |doi=10.1111/j.1469-185X.1987.tb01635.x |issn=1469-185X |s2cid=83672017}}</ref> 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.<ref name=":3">{{Cite journal |last=Ahlberg |first=Per Erik |date=1991-11-01 |title=A re-examination of sarcopterygian interrelationships, with special reference to the Porolepiformes |url=https://www.academia.edu/407873 |journal=Zoological Journal of the Linnean Society |language=en |volume=103 |issue=3 |pages=241–287 |doi=10.1111/j.1096-3642.1991.tb00905.x |issn=0024-4082}}</ref> 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.<ref name=":0">{{Cite journal |last=Jarvik |first=Erik |date=15 April 1996 |title=The Devonian tetrapod ''Ichthyostega'' |url=https://foreninger.uio.no/ngf/FOS/pdfs/F&S_40.pdf |journal=Fossils and Strata |volume=40 |pages=1–206|doi=10.18261/8200376605-1996-01 |isbn=82-00-37660-5 }}</ref><ref name=":4">{{Citation |last1=Borgen |first1=Ulf J. |title=Morphology, phylogeny and taxonomy of osteolepiform fish |date=2016 |pages=1–481 |publisher=Wiley-Blackwell |isbn=978-1-119-28643-1 |last2=Nakrem |first2=Hans A.|series=Fossils and Strata |volume=61 |doi=10.1002/9781119286448.ch1 }}</ref>
An alternative interpretation was proposed by T.S. Westoll (1938, 1943)<ref name=":5">{{Cite journal |last=Westoll |first=T. Stanley |date=1938 |title=Ancestry of the Tetrapods |url=https://www.nature.com/articles/141127a0 |journal=Nature |language=en |volume=141 |issue=3559 |pages=127–128 |doi=10.1038/141127a0 |bibcode=1938Natur.141..127W |s2cid=4086668 |issn=1476-4687|url-access=subscription }}</ref><ref name=":6">{{Cite journal |last=Westoll |first=T. S. |title=The Origin of the Tetrapods |date=1943 |url=https://onlinelibrary.wiley.com/doi/10.1111/j.1469-185X.1943.tb00289.x |journal=Biological Reviews |language=en |volume=18 |issue=2 |pages=78–98 |doi=10.1111/j.1469-185X.1943.tb00289.x |s2cid=85648376 |issn=1464-7931|url-access=subscription }}</ref> and A.S. Romer (1941).<ref>{{Cite journal |last=Romer |first=Alfred Sherwood |date=1941 |title=Notes on the crossopterygian hyomandibular and braincase |url=https://onlinelibrary.wiley.com/doi/10.1002/jmor.1050690106 |journal=Journal of Morphology |language=en |volume=69 |issue=1 |pages=141–160 |doi=10.1002/jmor.1050690106 |s2cid=85150870 |issn=0362-2525|url-access=subscription }}</ref> 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.<ref name=":5" /><ref name=":6" />
The Westoll/Romer hypothesis was initially controversial, with the strongest critiques coming from Scandinavian paleoichthyologists.<ref name=":7">{{Cite journal |last=Jarvik |first=E. |date=1967 |title=The homology of frontal and parietal bones in fishes and tetrapods |journal=Coll. Internatn. C.N.R.S. |volume=163 |pages=181–213}}</ref><ref name=":8">{{Cite journal |last=Borgen |first=Ulf J. |date=1983 |title=Homologizations of skull roofing bones between tetrapods and osteolepiform fishes |url=https://www.palass.org/sites/default/files/media/publications/palaeontology/volume_26/vol26_part4_pp735-753.pdf |journal=Palaeontology |volume=26 |issue=4 |pages=735–753}}</ref><ref name=":0" /><ref name=":4" /> 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.<ref name=":7" /><ref name=":8" /><ref name=":0" /><ref name=":4" /> Despite the criticism, the W/R scenario is now accepted by most paleontologists studying the fish-tetrapod transition.<ref>{{Cite journal |last=Parrington |first=F. R. |date=1967-10-01 |title=The identification of the dermal bones of the head |url=https://academic.oup.com/zoolinnean/article/47/311/231/2680225 |journal=Zoological Journal of the Linnean Society |language=en |volume=47 |issue=311 |pages=231–239 |doi=10.1111/j.1096-3642.1967.tb01406.x |issn=0024-4082|url-access=subscription }}</ref><ref>{{Cite journal |last1=Schultze |first1=H.P. |last2=Arsenault |first2=M. |date=1985 |title=The Panderichthyid fish Elpistostege: a close relative of tetrapods? |url=https://www.palass.org/sites/default/files/media/publications/palaeontology/volume_28/vol28_part2_pp293-309.pdf |journal=Palaeontology |volume=28 |issue=2 |pages=293–309}}</ref><ref name=":2" /><ref name=":3" /><ref name=":1" /> Developmental studies show support for the novel evolution of the frontal bones in tetrapods and their lack of homology with the "frontals" of fish.<ref>{{Cite journal |last1=Teng |first1=Camilla S |last2=Cavin |first2=Lionel |last3=Maxson |first3=Robert E Jnr |last4=Sánchez-Villagra |first4=Marcelo R |last5=Crump |first5=J Gage |date=2019-12-23 |editor-last=Tautz |editor-first=Diethard |title=Resolving homology in the face of shifting germ layer origins: Lessons from a major skull vault boundary |journal=eLife |volume=8 |article-number=e52814 |doi=10.7554/eLife.52814 |pmid=31869306 |pmc=6927740 |issn=2050-084X |doi-access=free }}</ref> Further tetrapodomorph discoveries, such as ''Acanthostega'' and ''Tiktaalik'',<ref name="Nature">{{cite journal |author=Edward B. Daeschler, Neil H. Shubin and Farish A. Jenkins Jr. |date=6 April 2006 |title=A Devonian tetrapod-like fish and the evolution of the tetrapod body plan |url=http://www.stuartsumida.com/BIOL524/DaeschlerEtAl2006.pdf |journal=Nature |volume=440 |issue=7085 |pages=757–763 |bibcode=2006Natur.440..757D |doi=10.1038/nature04639 |pmid=16598249 |doi-access=free}}</ref> provide more concrete evidence for the anatomical shifts hypothesized by Westoll and Romer.
==Diapsids and synapsids== [[File:SphenodonSkull.jpg|thumb|left|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: * The synapsids have one opening on each side, fairly low on the side of the skull, between the jugal or zygomatic bone and the elements above. Synapsids include mammals and their reptile-like ancestors (historically described under the misnomer "mammal-like reptiles"). In mammals, the side opening is closed by the sphenoid bone, so that the skull roof appears whole, despite the temporal opening. * The diapsids have two openings on each side, the two openings separated by an arch formed from processes of the postorbital and squamosal bones. All modern reptiles (apart from turtles) are diapsids in an anatomical sense, as are the birds and their dinosaur ancestors.
==References== {{Reflist}}{{Tetrapod osteology}}
Category:Skull Category:Vertebrate anatomy Category:Human head and neck Category:Flat bones