Placoderm

Placoderms (from Ancient Greek πλάξ [plax, plakos] 'plate' and δέρμα [derma] 'skin') are vertebrate animals of the class Placodermi, an extinct group of prehistoric fish known from Paleozoic fossils during the Silurian and the Devonian periods. While their endoskeletons are mainly cartilaginous, their head and thorax were covered by articulated armoured plates (hence the name), and the rest of the body was scaled or naked depending on the species.

Placoderms were among the first jawed fish (their jaws likely evolved from the first pair of gill arches), as well as the first vertebrates to have true teeth. They were also the first fish clade to develop pelvic fins, the second set of paired fins and the homologous precursor to hindlimbs in tetrapods. 380-million-year-old fossils of three other genera, Incisoscutum, Materpiscis and Austroptyctodus, represent the oldest known examples of live birth.

Placoderms are thought to be paraphyletic, consisting of several distinct outgroups or sister taxa to all living jawed vertebrates, which originated among their ranks. In contrast, one 2016 analysis concluded that Placodermi is likely monophyletic.

The first identifiable placoderms appear in the fossil record during the late Llandovery epoch of the early Silurian. They eventually outcompeted the previously dominant marine arthropods (e.g. eurypterids) and cephalopod molluscs (e.g. orthocones), producing some of the first and most infamous vertebrate apex predators such as Eastmanosteus, Dinichthys and the massive Dunkleosteus. Various groups of placoderms were diverse and abundant during the Devonian, but all placoderms became extinct at the end-Devonian Hangenberg event 358.9 million years ago, leaving the niches open for the osteichthyan and chondrichthyan survivors who subsequently radiated during the Carboniferous.

Many placoderms, particularly the Rhenanida, Petalichthyida, Phyllolepida, and Antiarchi, were bottom-dwellers. In particular, the antiarchs, with their highly modified, jointed bony pectoral fins, were highly successful inhabitants of Middle-Late Devonian freshwater and shallow marine habitats, with the Middle to Late Devonian genus, Bothriolepis, known from over 100 valid species. The vast majority of placoderms were predators, many of which lived at or near the substrate. Many, primarily the arthrodires, were active, nektonic predators that dwelled in the middle to upper portions of the water column. A study of the arthrodire Compagopiscis published in 2012 concluded that placoderms (at least this particular genus) likely possessed true teeth contrary to some early studies. The teeth had well defined pulp cavities and were made of both bone and dentine. However, the tooth and jaw development were not as closely integrated as in modern gnathostomes. These teeth were likely homologous to the teeth of other gnathostomes.

One of the largest known arthrodires, Dunkleosteus terrelli, was 3.5–4.1 metres (11–13 ft) long, and is presumed to have had a large distribution, as its remains have been found in Europe, North America and possibly Morocco. Some paleontologists regard it as the world's first vertebrate "superpredator", preying upon other predators. Other, smaller arthrodires, such as Fallacosteus and Rolfosteus, both of the Gogo Formation of Western Australia, had streamlined, bullet-shaped head armor, and Amazichthys, with morphology like that of other fast-swimming pelagic organisms, strongly supporting the idea that many, if not most, arthrodires were active swimmers, rather than passive ambush-hunters whose armor practically anchored them to the sea floor. Some placoderms were herbivorous, such as the Middle to Late Devonian arthrodire Holonema, and some were planktivores, such as the gigantic arthrodire Titanichthys, various members of Homostiidae, and Heterosteus.

Extraordinary evidence of internal fertilization in a placoderm was afforded by the discovery in the Gogo Formation, near Fitzroy Crossing, Kimberley, Western Australia, of a small female placoderm, about 25 cm (10 in) in length, which died in the process of giving birth to a 6 cm (2+1⁄2 in) offspring and was fossilized with the umbilical cord intact. The fossil, named Materpiscis attenboroughi (after scientist David Attenborough), had eggs which were fertilized internally, the mother providing nourishment to the embryo and giving birth to live young. With this discovery, the placoderm became the oldest vertebrate known to have given birth to live young ("viviparous"), pushing the date of first viviparity back some 200 million years earlier than had been previously known. Specimens of the arthrodire Incisoscutum ritchei, also from the Gogo Formation, have been found with embryos inside them indicating this group also had live bearing ability. The males reproduced by inserting a long clasper into the female. Elongated basipterygia are also found on the phyllolepid placoderms, such as Austrophyllolepis and Cowralepis, both from the Middle Devonian of Australia, suggesting that the basipterygia were used in copulation.

The placoderm claspers are not homologous with the claspers in cartilaginous fishes. The similarities between the structures has been revealed to be an example of convergent evolution. While the claspers in cartilaginous fishes are specialized parts of their paired pelvic fins that have been modified for copulation due to changes in the hox genes hoxd13, the origin of the mating organs in placoderms most likely relied on different sets of hox genes and were structures that developed further down the body as an extra and independent pair of appendages, but which during development turned into body parts used for reproduction only. Because they were not attached to the pelvic fins, as are the claspers in fish like sharks, they were much more flexible and could probably be rotated forward.