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Stegodon

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Stegodon

Stegodon (from the Ancient Greek στέγω (stégō), meaning "to cover", and ὀδούς (odoús), meaning "tooth", named for the distinctive ridges on the animal's molars) is an extinct genus of proboscidean, related to elephants. It was originally assigned to the family Elephantidae along with modern elephants but is now placed in the extinct family Stegodontidae. Like elephants, Stegodon had teeth with plate-like lophs that are different from those of more primitive proboscideans like gomphotheres and mammutids. Fossils of the genus are known from Africa and across much of Asia, as far southeast as Timor (with a single record in southeast Europe). The oldest fossils of the genus are found in Late Miocene strata in Asia, likely originating from the more archaic Stegolophodon, subsequently migrating into Africa. While the genus became extinct in Africa during the Pliocene, Stegodon persisted in South, Southeast and Eastern Asia into the Late Pleistocene.

The skull of Stegodon is relatively tall but short, and similar in many respects to those of living elephants. The lower jaw in comparison to early elephantimorphs and its ancestor Stegolophodon is shortened (brevirostrine), and lacks lower tusks/incisors. The molar teeth are superficially like those of elephants, consisting of parallel lamellae that form ridges but are generally relatively low crowned (brachydont), the numbers of ridges are greater in later species. Members of the genus lack permanent premolars. The tusks are proportionally large, with those of the biggest species being among the largest known tusks in proboscideans, with a particularly large tusk of S. ganesa from the Early Pleistocene of India measured to be 3.89 metres (12.8 ft) long, with an estimated mass of approximately 140 kilograms (310 lb), substantially larger than the largest recorded modern elephant tusk. These tusks have a slight upward curvature, and project forwards and parallel to each other, with the tusks often so close together that they are almost touching, such that the trunk would probably have had to rest on top of the tusks rather than be freely hanging between them as in living elephants.

The Chinese S. zdanskyi is suggested to be the largest species, and is known from an old male (50-plus years old) from the Yellow River that is 3.87 m (12.7 ft) tall and would have weighed approximately 12.7 tonnes (12.5 long tons; 14.0 short tons) in life. It had a humerus 1.21 m (4.0 ft) long, a femur 1.46 m (4.8 ft) long, and a pelvis 2 m (6.6 ft) wide. The Indian S. ganesa is suggested to have a shoulder height of about 3.10 m (10.2 ft), and a body mass of around 6.5 tonnes (6.4 long tons; 7.2 short tons). The Javanese species S. trigonocephalus is suggested to have been around 2.75–2.8 m (9.0–9.2 ft) tall, with a body mass of around 5 tonnes (4.9 long tons; 5.5 short tons). S. orientalis was around the size of an Asian elephant (Elephas maximus).

Similar to modern-day elephants, stegodonts were likely good swimmers, allowing them to disperse to remote islands in Indonesia, the Philippines and Japan. Once present on the islands, due to the process of insular dwarfism, as a result of decreased land area and the reduction of predation and competition pressure, they reduced in body size, with the degree of dwarfism varying between islands as the result of local conditions. One of the smallest species, Stegodon sumbaensis from Sumba in Indonesia, is estimated at around 8% of the size of mainland Stegodon species, with a body mass of 250 kilograms (550 lb). Sometimes the same island was colonised multiple times by Stegodon, as in Flores, where the Early Pleistocene strongly dwarfed species Stegodon sondaari, which was 120 cm (3.9 ft) tall at the shoulder and weighed about 350–400 kilograms (770–880 lb), was replaced by the species Stegodon florensis during the Middle Pleistocene which was initially substantially larger, but progressively reduced in size over time, with the earlier subspecies Stegodon florensis florensis from the Middle Pleistocene estimated to be around 50% the size of mainland Stegodon species with a shoulder height of around 190 cm (6.2 ft) and a body mass of around 1.7 tons, while the later Stegodon florensis insularis from the Late Pleistocene is estimated to be around 17% the size of mainland Stegodon species, with a shoulder height of around 130 cm (4.3 ft), and a body mass of about 570 kilograms (1,260 lb).

During Pliocene-Early Pleistocene (from around 4-1 million years ago), a succession of endemic dwarf species of Stegodon, probably representing a single lineage lived in the Japanese archipelago, probably derived from the mainland Chinese S. zdanskyi. In chronological succession these species are Stegodon miensis (4-3 million years ago) Stegodon protoaurorae (3-2 million years ago) and Stegodon aurorae, (2-1 million years ago) which show a progressive size reduction through time, possibly as a result of reducing land area of the Japanese archipelago. The latest and smallest species S. aurorae is estimated to be 25% the size of its mainland ancestor with a body mass of around 2,122 kilograms (4,678 lb). S. aurorae also shows morphological straits associated with dwarfism, like shortened limbs.

Like modern elephants, but unlike more primitive proboscideans, Stegodon is thought to have chewed using a proal movement (a forward stroke from the back to the front) of the lower jaws. This jaw movement is thought to have evolved independently in elephants and stegodontids. Stegodon populations from the Late Pliocene of the India (including Stegodon insignis) are suggested to have been variable mixed feeders, while those from the earliest Pleistocene (including Stegodon ganesa) of the same region are suggested to have been nearly pure grazers based on isotopic analysis. Based on dental microwear analysis, populations of Stegodon from the Pleistocene of China (Stegodon orientalis and Stegodon huananensis) and mainland southeast Asia (S. orientalis) were found to be browsers, with clear niche differentiation from sympatric Elephas populations, which tended towards mixed feeding (both browsing and grazing), though isotopic analysis of Stegodon cf. orientalis specimens from the late Middle Pleistocene of Thailand suggests that these individuals were mixed feeders that consumed a significant amount of C4 grass. Specimens of Stegodon trigonocephalus from the Early-Middle Pleistocene of Java were found to be mixed feeders to grazers, with a diet similar to that of sympatric Elephas hysudrindicus. The dwarf species from Flores, Stegodon sondaari and Stegodon florensis, are suggested to have been mixed feeders and grazers, respectively, based on stable carbon isotopes. Specimens of Stegodon kaiesensis from the Pliocene of East Africa were found to be browsers to mixed feeders, based on mesowear analysis.

Tracks of a group of Stegodon from the Late Pliocene of Japan suggest that like modern elephants, Stegodon lived in social herds.

On Flores, where dwarf Stegodon species were the only large herbivores, they were likely the main prey of the Komodo dragon.

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