Acanthodii

"Spiny shark" redirects here. For the spiny true sharks, see Echinorhinus.

Acanthodii Temporal range: Early Silurian–Permian PreЄ Є O S D C P T J K Pg N
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Clade:	Craniata
Subphylum:	Vertebrata
Infraphylum:	Gnathostomata
Class:	†Acanthodii Owen, 1846
Orders
Climatiiformes Ischnacanthiformes Acanthodiformes

Acanthodii (sometimes called spiny sharks) is a class of extinct fishes, sharing features with both bony fish and cartilaginous fish. In form they resembled sharks, but their epidermis was covered with tiny rhomboid platelets like the scales of holosteans (gars, bowfins). They may have been an independent phylogenetic branch of fishes, which had evolved from generalized or basal forms similar to primitive members of the still-extant Chondrichthyes.

The popular name "spiny sharks" is really a misnomer for these early jawed fishes. The name was coined because they were superficially shark-shaped, with a streamlined body, paired fins, and a strongly upturned tail; stout, largely immovable bony spines supporting all the fins except the tail - hence, "spiny sharks". Fossilized spines and scales are often all that remains of these fishes in ancient sedimentary rocks. Although not sharks or cartilaginous fish, acanthodians did, in fact, have a cartilaginous skeleton, but their fins had a wide, bony base and were reinforced on their anterior margin with a dentine spine. The earliest acanthodians were marine, but during the Devonian, freshwater species became predominant.

There are three orders recognized: Climatiiformes, Ischnacanthiformes and Acanthodiformes. Climatiiforma had shoulder armor and many small sharp spines, Ischnacanthiforma with teeth fused to the jaw, and the Acanthodiforma were filter feeders, with no teeth in the jaw, but long gill rakers. Overall, the acanthodians' jaws are presumed to have evolved from the first gill arch of some ancestral jawless fishes that had a gill skeleton made of pieces of jointed cartilage.

Characteristics

Mesacanthus, Parexus, and Ischnacanthus of Early Devonian Great Britain

Diplacanthus longispinus impression at the Museum für Naturkunde, Berlin

The scales of Acanthodii have distinctive ornamentation peculiar to each order. Because of this, the scales are often used in determining relative age of sedimentary rock. The scales are tiny, with a bulbous base, a neck, and a flat or slightly curved diamond-shaped crown.

Despite being called "spiny sharks," acanthodians predate sharks. Scales that have been tentatively identified as belonging to acanthodians, or "shark-like fishes" have been found in various Ordovician strata, though, they are ambiguous, and may actually belong to jawless fishes such as thelodonts. The earliest unequivocal acanthodian fossils date from the beginning of the Silurian Period, some 50 million years before the first sharks appeared. Later, the acanthodians colonized fresh waters, and thrived in the rivers and lakes during the Devonian and in the coal swamps of Carboniferous. But the first bony fishes were already showing their potential to dominate the waters of the world, and their competition proved too much for the spiny sharks, which died out in Permian times (approximately 250 MYA).

Many paleonthologists consider that the acanthodians were close to the ancestors of the bony fishes. Although their interior skeletons were made of cartilage, a bonelike material had developed in the skins of these fishes, in the form of closely fitting scales (see above). Some scales were greatly enlarged and formed a bony covering on top of the head and over the lower shoulder girdle. Others developed a bony flap over the gill openings analogous to the operculum in later bony fishes.

Taxonomy and phylogeny

In a study of early jawed vertebrate relationships, Davis et al. (2012) found acanthodians to be split among the two major clades Osteichthyes (bony fish) and Chondrichthyes (cartilaginous fish). The well-known acanthodian Acanthodes was placed within Osteichthyes, despite the presence of many chondrichthyan characteristics in its braincase. Below is a cladogram from the analysis, with bold lines representing the evolutionary grade of acanthodians:

Vertebrata  Galeaspida Osteostraci Gnathostomata Bothriolepis  Pterichthyodes  Brindabellaspis  Macropetalichthys  Dicksonosteus  Buchanosteus  Coccosteus  Cowralepis  Austroptyctodus  Campbellodus  Rhamphodopsis  Tetanopsyrus  Culmacanthus  Diplacanthus  Gladiobranchus  Rhadinacanthus  Ptomacanthus  Climatius  Brachyacanthus  Parexus  Osteichthyes Euthacanthus  Cassidiceps  Mesacanthus  Promesacanthus  Cheiracanthus  Acanthodes  Homalacanthus  Ischnacanthus  Poracanthodes  Dialipina Ligulalepis Crown group Osteichthyes Chondrichthyes Vernicomacanthus  Brochoadmones  Kathemacanthus  Obtusacanthus  Lupopsyrus  Pucapampella Doliodus Tamiobatis Cladodoides Orthacanthus Cobelodus Akmonistion Cladoselache Tristychius Crown group Chondrichthyes

Placodermi Acanthodii Acanthodii

See also

• List of acanthodians
• Placoderm

Further reading

• Beneš, Josef (1979), Prehistoric Animals and Plants, New York: Hamlyn, ISBN 0-600-30341-1 
• Janvier, Phillipe (1996), Early vertebrates, Oxford: Clarendon Press, ISBN 0-19-854047-7 
• Long, John A. (1995), The Rise of Fishes: 500 Million Years of Evolution, Baltimore: Johns Hopkins University Press, ISBN 0-8018-4992-6 
• Palmer, Douglas, ed. (1999), The Simon & Schuster Encyclopedia of Dinosaurs & Prehistoric Creatures. A Visual Who's Who of Prehistoric Life, New York: Simon & Schuster, ISBN 0-684-86411-8 

External links

• Acanthodii taxonomy
• †Ischnacanthiformes taxonomy
• †Climatiiformes taxonomy
• †Acanthodiformes taxonomy
• Ischnacanthus gracilis - A Devonian Fish by Neal Robbins
• PALAEOZOIC FISH UK
• Acanthodopsis wardi