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Gorgonopsia

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Gorgonopsia

Gorgonopsia (from the Greek Gorgon, a mythological beast, and óps 'aspect') is an extinct clade of sabre-toothed therapsids from the Middle to the Upper Permian, roughly between 270 and 252 million years ago. They are characterised by a long and narrow skull, as well as elongated upper and sometimes lower canine teeth and incisors which were likely used as slashing and stabbing weapons. Postcanine teeth are generally reduced or absent. For hunting large prey, they possibly used a bite-and-retreat tactic, ambushing and taking a debilitating bite out of the target, and following it at a safe distance before its injuries exhausted it, whereupon the gorgonopsian would grapple the animal and deliver a killing bite. They would have had an exorbitant gape, possibly in excess of 90°, without having to unhinge the jaw.

They markedly increased in size as time went on, growing from small skull lengths of 10–15 cm (4–6 in) in the Middle Permian to bear-like proportions of up to 60 cm (2 ft) in the Upper Permian. The latest gorgonopsians, Rubidgeinae, were the most robust of the group and could produce especially powerful bites. Gorgonopsians are thought to have been completely terrestrial and could walk with a semi-erect gait, with a similar terrestrial locomotory range as modern crocodilians. They may have been more agile than their prey items, but were probably inertial homeotherms rather than endotherms unlike contemporary therocephalians and cynodonts, and thus were probably comparatively less active. Though gorgonopsians were able to maintain a rather high body temperature, it is unclear if they would have also had sweat glands or fur (and by extension whiskers and related structures). Their brains were reminiscent of modern reptilian brains, rather than those of living mammals. Most species may have been predominantly diurnal (active during the day) though some could have been crepuscular (active at dawn or dusk) or nocturnal (active at night). They are thought to have had binocular vision, a parietal eye (which detects sunlight and maintains circadian rhythm), a keen sense of smell, a functional vomeronasal organ ("Jacobson's organ"), and possibly a rudimentary eardrum.

The major therapsid groups had all evolved by 275 million years ago from a "pelycosaur" ancestor (a poorly defined group including all synapsids which are not therapsids). The therapsid takeover from pelycosaurs took place by the Middle Permian as the world progressively became drier. Gorgonopsians rose to become apex predators of their environments following the Capitanian mass extinction event which killed off the dinocephalians and some large therocephalians after the Middle Permian. Despite the existence of a single continent during the Permian, Pangaea, gorgonopsians have only been found in the Karoo Supergroup (primarily in South Africa, but also in Tanzania, Zambia, and Malawi), the Moradi Formation of Niger, western Russia, the Port des Canonge Formation on the island of Mallorca in Spain, and in the Turpan Basin of Xinjiang, China, with probable remains known from the Kundaram Formation in the Pranhita–Godavari Basin of India. These places were semi-arid areas with highly seasonal rainfall. The oldest gorgonopsian specimen, a middle Permian gorgonopsian, was described from the Port des Canonge Formation on the island of Mallorca, western Mediterranean. Gorgonopsian genera are all very similar in appearance, and consequently many species have been named based on flimsy and likely age-related differences since their discovery in the late 19th century, and the group has been subject to several taxonomic revisions.

Most gorgonopsians became extinct during a phase of the Permian–Triassic extinction event taking place at the very end of the Permian, in which major volcanic activity (which would produce the Siberian Traps) and resultant massive spike in greenhouse gases caused rapid aridification due to temperature spike, acid rain, frequent wildfires, and potential breakdown of the ozone layer. However, some smaller taxa like Cyonosaurus may have survived up to the Early Triassic. The large predatory niches would be taken over by the archosaurs (namely crocodilians and dinosaurs) in the Mesozoic.

Earlier gorgonopsids in the Middle Permian were quite small, with skull lengths of 10–15 cm (4–6 in), whereas some later genera attained massive, bear-like sizes with the largest being Inostrancevia up to 3.5 m (11 ft) in length and 300 kg (660 lb) in body mass. Nonetheless, small gorgonopsians remained abundant until extinction (though small species may actually represent juvenile specimens of other taxa).

Like other Permian therapsids, gorgonopsians had developed several mammalian characteristics. These might have included a parasagittal gait (the limbs were vertically oriented and moved parallel to the spine) as opposed to the sprawling gait of amphibians and earlier synapsids. This gait change in therapsids was possibly related to the reduction in tail size and phalangeal formula (the number of bones per digit, which for gorgonopsians was 2.3.4.5.3 like reptiles). Other developments included fibrous lamellar cortical bone and deeply-set teeth. Like reptiles, gorgonopsians lack a secondary palate separating the mouth from the nasal cavity, prohibiting chewing.

Anatomy varies incredibly little between gorgonopsians. Many species are distinguished by vague proportional differences, and consequently smaller species may actually represent juveniles of larger taxa. Notably, the vomer at the tip of the snout varies among species in terms of the degree of its expansion, as well as the positions, degree of splay, and shape of the 3 ridges. They typically feature a long and narrow skull. Juvenile Rubidgea appear to have had snouts wider than long. Unlike eutheriodonts, the occipital bone (at the back of the skull) is rectangular (wider than tall) and concave, as opposed to triangular.

The gorgonopsian brain, like other non-mammaliaform therapsids, lacks an expansion of the neocortex, has a relatively large hindbrain compared to the forebrain, a large epyphysial nerve (found in creatures with a parietal eye on the top of the head), an enlarged pituitary gland, and an overall elongated shape; all-in-all resembling a reptilian brain. The braincase was also rather reptilian, and is also comparatively smaller and not as thick as those of mammals. The flocculus, a lobe of the cerebellum, is proportionally large, and is related to the vestibulo-ocular reflex (which stabilises gaze while moving the head). Judging by the orientation of the semi-circular canals in the ear (which have to be oriented parallel to the ground), the head of the gorgonopsian specimen GPIT/RE/7124 would have tilted forward by about 41°, increasing the overlap between the visual fields of the two eyes and improving binocular vision—useful to a predator. Unlike either reptiles or mammals, the semi-circular canals are flat, probably because they were wedged between the opisthotic (an inner ear bone) and supraoccipital bones.

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