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Sarcopterygian Fishes, the “Lobe-Fins”

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Heads, Jaws, and Muscles

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Abstract

Sarcopterygian fishes, the “lobe-fins”, are today represented by two species of coelacanth and six lungfishes. Lungfish are the closest living group to the tetrapods, the first four-footed terrestrial vertebrates and all of their descendants. However, when extinct groups are taken into account, the evolutionary history and relationships of the Sarcopterygii become much more complex. In addition to lungfishes and the coelacanth, this chapter will introduce you to groups now known exclusively from fossils such as “dagger-toothed” onychodonts and porolepiforms named so for the special pores in their scales. Furthermore, the fish-tetrapod transition occurred gradually with many stem-tetrapods (finned tetrapods) progressively acquiring characters that would later become fixed features of terrestrial vertebrates. Primitive sarcopterygians have skulls that are divided in two halves, a feature still observable in the coelacanth today but lost in lungfishes and tetrapods. Lobe-finned fishes have teeth made of dentine and enamel; early members also possessed a tissue called cosmine covering their skulls and scales. A large variety of dentitions are visible across the Sarcopterygii, such as the fearsome tooth whorls in onychodonts and porolepiforms, crushing tooth plates in lungfishes or dagger-like fangs in tetrapodomorphs. The first appearance of an inner ear bone (stapes), an internal nostril (choana) and separation of the skull and shoulder girdle to form a neck occurred within this group. The earliest tetrapodomorphs retained fins and were still fully aquatic and are known from the Early Devonian onwards.

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Acknowledgments

Firstly I wish to thank John Long, Brian Choo and Min Zhu for their generous supply of images included in this chapter and Jing Lu for her helpful comments on the text of an earlier draft. I would also like to thank the editors, Janine Ziermann, Raul Diaz Jr, and Rui Diogo for the invitation to contribute this chapter. Two helpful reviews from John Long and Sébastian Olive improved the quality of the text.

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Correspondence to Alice M. Clement .

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Glossary

Acanthodian/s

A group of stem-chondrichthyans known as “spiny sharks” with bony spines preceding all their fins

Actinistia

A group of sarcopterygian fish commonly known as coelacanths

Actinopterygian/s

One of the major groups of bony fishes, comprising the vast majority of extant fish species, also known as “ray-finned fishes”

Adductor mandibulae/mandibular/muscle

The major jaw muscles

Aestivate

To spend a long period in torpor during hot or dry conditions (similar to hibernation)

Amniota

A group of vertebrates that can lay eggs with an amnion (a membrane that covers the embryo), this group includes all reptiles, birds and mammals

Ampullae of Lorenzini

Special electroreceptive sense organs (jelly-filled pores) found commonly in cartilaginous fish (e.g., sharks, rays and chimaeras)

Articular bone

A bone that is part of the lower jaw in most vertebrates

Autostyly/autostylic jaw

A type of jaw suspension whereby the upper jaw is connected directly to the cranium

Basicranial muscle

Large muscles that span the intracranial joint and attach underneath the braincase that is activated during feeding

Braincase

Also known as the “neurocranium”, this is the inner part of the skull that encases the brain

Branchial

Meaning related to the gills

Carboniferous

A geological period that occurred 299–359 million years ago

Cartilaginous

A type of connective tissue found in the body, it can be mineralized and form part of the skeleton, but it is not as hard as bone, but far stiffer than muscle

Cenozoic

The geological era that occurred from 23 million years ago to the present

Choana

A unique internal nostril found in tetrapodomorphs which opens from the nasal sac into the roof of the mouth

Chondrichthyans

A group of fish with their skeletons made primarily of cartilage rather than bone (e.g., sharks, rays and chimaeras)

Class

A taxonomic rank in biological classification (e.g., Mammalia)

Cleavage

Division of the cells in the early embryo

Convergent

Independent evolution of similar features in different lineages

Coracomandibularis

A muscle found in coelacanths that elevates the palatoquadrate

Coronoid

A bone forming part of the lower jaw

Cosmine

A hard tissue present on the scales and dermal bones in primitive sarcopterygians that is composed of a mixture of enamel and dentine

Cranial

A subdivision of the skull (that together with the mandible comprises the skull)

Cranial ribs

Paired structures that attach to the base of the braincase in some lungfishes, distinct from the pleural ribs

Cretaceous

A geological period that occurred 66–145 million years ago

Cristae

Bony struts supporting the skull roof in primitive lungfishes, allowing large spaces for attachment of jaw muscles

Dendrodont

A special type of dentition found in porolepiforms whereby their fangs show a infolding of enamel and dentine

Dentary

The main dermal bone of the lower jaw

Denticulated

A type of dentition seen in some early lungfishes whereby their buccal cavity is covered with a shagreen of small denticles (e.g., Griphognathus)

Dentine

A calcified tissue of the body, one of the major components of teeth

Dentine-plated

A type of dentition seen in some early lungfishes whereby large areas within their buccal cavity are covered in very hard but usually smooth hypermineralised dentine patches

Dentition

Arrangement or condition of teeth

Depressor mandibulae

A muscle used to open the jaw (in lungfishes)

Dermal skull

The skull roof, or roofing bones of the skull derived from dermal bone

Devonian

A geological period that occurred 359–419 million years ago

Dipnoi

A group of sarcopterygian fish commonly known as lungfishes

Durophagy

Feeding by crushing prey, usually hard-shelled animals

Electroreception/electrosensory

The biological ability to perceive electrical stimuli

Elpistostegalids

A group of advanced stem-tetrapods such as Tiktaalik

Enamel

The hardest type of calcified tissue of the body, one of the major components of teeth

Endocast

Internal cast of a hollow object, such as the cavity inside the skull

Epaxials

Dorsal trunk body muscles

Ethmosphenoid

The front portion of the skull

Extant

Living or recent, not extinct

Frontals

Paired median skull roof bones at the front of the skull in tetrapod-related taxa

Ganoine

A bony tissue homologous to enamel that is found in actinopterygians

Gastrulation

An early phase of embryonic development

Geniocoracoideus

A muscle opening jaw found in the lungfish Neoceratodus

Geniothoracicus

A muscle opening jaw found in the lepidosirenid lungfishes

Genome

The full genetic material of an organism

Gnathostome

Jawed vertebrates

Gogo Formation

A geological formation and famous Devonian fossil site in North Western Australia

Homologies/homologous/homology

Shared ancestry between structures in different taxa

Hyoid

A bone derived from the second gill arch in fish, used during feeding and respiration

Hyomandibula

One of the jaw attachment bones in fishes that become incorporated into the inner ear of tetrapods

Hypaxials

Ventral trunk body muscles

Hypermineralised

Highly mineralized

Hypobranchial

Meaning located below the gills

In vivo

Within a living organism

Inner ear

The portion of the ear located within the skull

Interhyoideus

A muscle used to help raise the jaw and hyoid in lungfishes

Intermandibularis

A muscle used to help raise the jaw and hyoid in lungfishes

Intracranial joint

The joint separating two divisions of the skull in early sarcopterygians and still present in the coelacanth

Kinetic/kinesis

Moveable

Labyrinthodont

Teeth with a complex pattern of infolding of dentine and enamel found in some tetrapods

Lateral line

A sense organ of fishes running the length of their body to detect vibration and movement in the surrounding water

Lepidosirenid

A member of the lepidosirenid family of lungfishes (Lepidosiren and Protopterus, but not Neoceratodus)

Levator hyoideus

A muscle used to help raise the jaw and hyoid in lungfishes

Lip retractor

Muscles used to control the lips in lepidosirenid lungfishes

Lissamphibia

A taxonomic group that includes all modern (crown group) amphibians but excludes stem members

Mandible

Lower jaw

Maxilla

An upper jaw bone

Mesozoic Era

A geological Era that occurred 66–252 million years ago

Metamorphosis

A biological process whereby an animal’s body undergoes conspicuous change and development after birth

Monophyletic

A cladistic term used to characterise a clade of organisms that share derived characters (synapomoprhies)

Neural crest

A temporary group of cells that give rise to a diverse range of cells in the body, unique to vertebrates

Onychodont

A group of extinct basal sarcopterygian fish related to Coelacanths

Opercular bones

A bony flap covering the gills in fishes

Ossified/ossifications

Meaning bony/bones

Osteichthyans

The “bony fishes” are a superclass that includes all Actinopterygii and Sarcopterygii

Osteolepiformes

A paraphyletic assemblage of finned stem-tetrapods from the Devonian Period (e.g., Eusthenopteron)

Otoccipital

The rear portion of the skull

Ovoviviparous

Live-bearing, embryos develop inside eggs that remain within the mother’s body

Paedomorphosis

The retention of juvenile traits into adulthood, or “juvenilisation”, a type of heterochrony

Palaeozoic Era

A geological Era that occurred 252–541 million years ago

Palatoquadrate

Endoskeletal part of the upper jaw

Parasphenoid

A median (often with teeth) bone of the palate in fishes

Parietal shield

A set of dermal bones covering the dorsal surface of the anterior half of the skull

Parietals

Paired skull bones that enclose the pineal region in fishes and tetrapods

Pectoral girdle

Shoulder girdle

Permian

The geological period that occurred 252–299 million years ago

Phylogenetic position/phylogeny

Inferred evolutionary relationships among taxa based on differences and similarities in their physical or genetic characteristics

Placoderm

An extinct group of fishes that were dominant during the Devonian with thick plated “armour” covering their bodies

Pleural

Referring to the body, especially around the chest cavity

Porolepiformes

A group of extinct sarcopterygian fishes closely related to lungfishes

Postcranial

All parts of the skeleton apart from the skull

Postparietals

Paired median skull roof bones situated towards the rear

Premaxilla

An upper jaw bone situated in front of the maxilla

Preopercular

A dermal cheekbone situated in front of the operculum

Preorbital

The area of a skull situated in front of the eyes (orbits)

Quadrate

A bone in the skull that contributes to the jaw joint

Quadratojugal

A cheekbone found in some fishes

Rectus cervicis

A muscle used to open the jaw (in lungfishes)

Rhizodonts

A monophyletic group of stem-tetrapods that reached huge sizes during the Palaeozoic

Rostral organ

A large sensory organ in the snout of coelacanths

Rostral tubuli

Network of small, bony tubules throughout the snout in some sarcopterygian fishes

Sarcopterygian/Sarcopterygii

One of the major groups of bony fishes that include lungfish, coelacanths and tetrapods, also known as “lobe-finned fishes”

Silurian

A geological period that occurred 419–444 million years ago

Sister taxa/group

The closest relatives of another taxon/group in a phylogenetic tree

Skull roof

The roofing bones of the skull derived from dermal bone

Spiracular openings/slits

Openings on the top of the skull in some fishes and tetrapods, thought to be involved in accessory air breathing

Squamosal

A cheekbone found in some fishes

Stapes

A bone of the inner ear found in tetrapods

Stem group

A phylogenetic term meaning members of a total group that are excluded from the crown group

Sutures

A type of fibrous joint between bones of the skull

Tandem jaw joint

A specialised jaw arrangement found in coelacanths whereby the lower jaw is attached via two joints on each side of the head

Terrestrial

Live predominantly on land (rather than in water)

Tetrapodomorph/s

Meaning “four-footed-like” and this includes finned and limbed forms that are more closely related to living tetrapods than to living lungfishes

Tetrapods

“Four-footed” vertebrates with digit-bearing limbs and all of their descendants

Tooth whorl

A distinctive type of dentition found in some sarcopterygian fishes

Tooth-plated

Those lungfishes with tooth plates rather than denticulated or dentine-plated dentition

Topologically

Pertaining to position or region

Urodeles

A group of amphibians that includes salamanders

Vascular

Carrying blood within the body, part of the circulatory system

Vertebrates

All animals with a backbone (including fishes, amphibians, reptiles, birds and mammals)

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Clement, A.M. (2019). Sarcopterygian Fishes, the “Lobe-Fins”. In: Ziermann, J., Diaz Jr, R., Diogo, R. (eds) Heads, Jaws, and Muscles. Fascinating Life Sciences. Springer, Cham. https://doi.org/10.1007/978-3-319-93560-7_6

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