Stegodon

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Stegodon
Temporal range: Late Miocene–Late Pleistocene, 11.6–0.01  Ma
Stegodon hunghoensis.JPG
Stegodon skeleton at the Gansu Provincial Museum
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Proboscidea
Family:† Stegodontidae
Genus:† Stegodon
Falconer, 1847
Species
  • † S. aurorae (Matsumoto, 1918)
  • †S. elephantoides(Clift, 1828)
  • †S. florensisHooijer, 1957
  • †S. ganesha
    (Faloner and Cautley, 1846)
  • †S. kaisensisHopwood, 1939
  • †S. luzonensis
    von Koenigswald, 1956
  • †S. miensis(Matsumoto, 1941)
  • †S. mindanensis(Naumann, 1890)
  • †S. orientalisOwen, 1870
  • † S. protoaurorae (Aiba et al., 2010)
  • †S. sompoensisHooijer, 1964
  • †S. sondaarivan den Bergh, 1999
  • †S. trigonocephalus(Martin, 1887)
  • †S. zdanskyiHopwood, 1935

Stegodon ("roofed tooth" from the Ancient Greek words στέγω , stégō, 'to cover', + ὀδούς , odoús, 'tooth' because of 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. [1] 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. [2] While the genus became extinct in Africa during the Pliocene, Stegodon remained widespread in South, Southeast and East Asia until the end of the Pleistocene. [3]

Morphology

The skull of Stegodon is relatively tall but short. [1] The lower jaw in comparison to early elephantimorphs 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), [1] [4] the numbers of ridges are greater in later species. [5] Members of the genus lack permanent premolars. [6] 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. [7]

Size

Skeletal restoration of a large male S. zdanskyi with a shoulder height of around 3.87 metres Stegodon skeletal.png
Skeletal restoration of a large male S. zdanskyi with a shoulder height of around 3.87 metres

Some species of Stegodon were amongst the largest proboscideans. The Chinese species S. zdanskyi 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). [8]

Dwarfism

Estimated sizes of dwarf Stegodon species from Flores compared to a human Dwarf stegodon size comparison.svg
Estimated sizes of dwarf Stegodon species from Flores compared to a human

Similar to modern-day elephants, stegodonts were likely good swimmers, [9] [10] 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). [11] 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), [8] 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). [11] [12]

Ecology

Life-sized models of Stegodon ExtinctElephant-SiwalikHills.JPG
Life-sized models of Stegodon

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. [1] Stegodon populations from the Pliocene of the India are suggested to have been variable mixed feeders based on isotopic analysis. [13] 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). [14] [15] In contrast, 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 . [16] 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. [12] Specimens of Stegodon kaiesensis from the Pliocene of East Africa were found to be browsers to mixed feeders, based on mesowear analysis. [17]

Tracks of a group of Stegodon from the Late Pliocene of Japan suggest that like modern elephants, Stegodon lived in social herds. [18]

On Flores, where dwarf Stegodon species were the only large herbivores, they were likely the main prey of the Komodo dragon. [19] Claims that Stegodon florensis was hunted by Homo floresiensis are based on ambiguous circumstantial association between bones and stone tools, and the rarity of cut marks makes it unclear to what if to any degree, hunting of Stegodon was actually practiced by Homo floresiensis. [20] [21]

Taxonomy

Fossils of S. aurorae (left) and S. orientalis (right) at the National Museum of Nature and Science, Tokyo Stegodon aurorae and Stegodon orientalis.jpg
Fossils of S. aurorae (left) and S. orientalis (right) at the National Museum of Nature and Science, Tokyo
Jaw fossil of S. sondaari at the Bandung Geological Museum Stegodon sondaari.jpg
Jaw fossil of S. sondaari at the Bandung Geological Museum
Skull of S. ganesha Stegodon ganesa.JPG
Skull of S. ganesha

In the past, stegodonts were believed to be the ancestors of the true elephants and mammoths, but currently they are believed to have no modern descendants. Stegodon is likely derived from Stegolophodon , an extinct genus known from the Miocene of Asia, [2] with transitional fossils between the two genera known from the Late Miocene of Southeast Asia and Yunnan in South China. [1] Stegodon is more closely related to elephants and mammoths than to mastodons. [22] Like elephants, stegodontids are believed to have derived from gomphotheres. [23]

Phylogeny

The following cladogram shows the placement of the genus Stegodon among other proboscideans, based on hyoid characteristics: [22]

Mammut americanum (American mastodon)

Gomphotherium sp.

Stegodon zdanskyi

Loxodonta africana (African bush elephant)

Elephas maximus (Asian elephant)

Mammuthus columbi (Columbian mammoth)

List of species

An indeterminate Stegodon molar of an uncertain locality and age is known from Greece, representing the only record of the genus in Europe. [24] Indeterminate remains are also known from the Early Pleistocene and early Middle Pleistocene of Israel. [25]

Evolution and extinction

The oldest fossils of Stegodon in Asia date to the Late Miocene, around 8-11 million years ago, [2] with the oldest fossils of the genus in Africa being around 7-6 million years old. [26] Stegodon became extinct in Africa during the late Pliocene, around 3 million years ago suggested to be the result of expansion of grassland habitats. [17] The Javanese species Stegodon trigonocephalus became extinct around 130-80,000 years ago during the latest Middle Pleistocene-early Late Pleistocene (Marine Isotope Stage 5) following a change to more humid conditions, which may have reduced grazing habitat. [16] Stegodon florensis became extinct around 50,000 years ago, around the time of arrival of modern humans to Flores. [27] Stegodon became extinct in the Indian subcontinent (Stegodon namadicus/Stegodon sp.), mainland Southeast Asia and China (S. orientalis) at some point during the Late Pleistocene epoch, while Asian elephants, which existed in sympatry with Stegodon in these regions, are still extant. The precise timing of extinction is uncertain for these regions. [28] [15] [29] The survival of the Asian elephant as opposed to Stegodon orientalis in Southeast Asia and South China has been suggested to be due to its more flexible diet in comparison to S. orientalis. [15] A review of 130 papers written about 180 different sites with proboscidean remains in southern China revealed Stegodon to have been more common than Asian elephants; the papers gave many recent radiocarbon dates, the youngest being 2,150 BCE (4,100 BP). [30] However, Turvey et al. (2013) reported that one of the faunal assemblages including supposed fossils of Holocene Stegodon (from Gulin, Sichuan Province) is actually late Pleistocene in age; other supposed fossils of Holocene stegodonts were lost and their age cannot be verified. The authors concluded that the latest confirmed occurrences of Stegodon from China are from the Late Pleistocene, and that its Holocene survival cannot be substantiated. [29]

Related Research Articles

<span class="mw-page-title-main">Mammoth</span> Extinct genus of mammals

A mammoth is any species of the extinct elephantid genus Mammuthus. They lived from the late Miocene epoch into the Holocene about 4,000 years ago, and various species existed in Africa, Europe, Asia, and North America. Mammoths are more closely related to living Asian elephants than African elephants.

<span class="mw-page-title-main">Proboscidea</span> Order of elephant-like mammals

Proboscidea is a taxonomic order of afrotherian mammals containing one living family (Elephantidae) and several extinct families. First described by J. Illiger in 1811, it encompasses the elephants and their close relatives. Three species of elephant are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant.

<span class="mw-page-title-main">Elephantidae</span> Family of mammals

Elephantidae is a family of large, herbivorous proboscidean mammals collectively called elephants and mammoths. These are large terrestrial mammals with a snout modified into a trunk and teeth modified into tusks. Most genera and species in the family are extinct. Only two genera, Loxodonta and Elephas, are living.

<span class="mw-page-title-main">Mammutidae</span> Extinct family of mammals

Mammutidae is an extinct family of proboscideans belonging to Elephantimorpha. It is best known for the mastodons, which inhabited North America from the Late Miocene until their extinction at beginning of the Holocene, around 11,000 years ago. The earliest fossils of the group are known from the Late Oligocene of Africa, around 24 million years ago, and fossils of the group have also been found across Eurasia. The name "mastodon" derives from Greek, μαστός "nipple" and ὀδούς "tooth", referring to their characteristic teeth.

<i>Elephas</i> Genus of mammals

Elephas is one of two surviving genera in the family of elephants, Elephantidae, with one surviving species, the Asian elephant, Elephas maximus. Several extinct species have been identified as belonging to the genus, extending back to the Pliocene era.

<i>Palaeoloxodon</i> Genus of extinct elephants

Palaeoloxodon is an extinct genus of elephant. The genus originated in Africa during the Early Pleistocene, and expanded into Eurasia at the beginning of the Middle Pleistocene. The genus contains some of the largest known species of elephants, over 4 metres (13 ft) tall at the shoulders, including the African Palaeoloxodon recki, the European straight-tusked elephant and the South Asian Palaeoloxodon namadicus. P. namadicus has been suggested to be the largest known land mammal by some authors based on extrapolation from fragmentary remains, though these estimates are highly speculative. In contrast, the genus also contains many species of dwarf elephants that evolved via insular dwarfism on islands in the Mediterranean, some only 1 metre (3.3 ft) in height, making them the smallest elephants known. The genus has a long and complex taxonomic history, and at various times, it has been considered to belong to Loxodonta or Elephas, but today is usually considered a valid and separate genus in its own right.

<span class="mw-page-title-main">Gomphothere</span> Extinct family of proboscidean mammals

Gomphotheres are an extinct group of proboscideans related to modern elephants. They were widespread across Afro-Eurasia and North America during the Miocene and Pliocene epochs and dispersed into South America during the Pleistocene as part of the Great American Interchange. Gomphotheres are a paraphyletic group that is ancestral to Elephantidae, which contains modern elephants, as well as Stegodontidae. While most famous forms such as Gomphotherium had long lower jaws with tusks, which is the ancestral condition for the group, some later members developed shortened (brevirostrine) lower jaws with either vestigial or no lower tusks, looking very similar to modern elephants, an example of parallel evolution, which outlasted the long-jawed gomphotheres. By the end of the Early Pleistocene, gomphotheres became extinct in Afro-Eurasia, with the last two genera, Cuvieronius ranging from southern North America to western South America, and Notiomastodon having a wide range over most of South America until the end of the Pleistocene around 12,000 years ago, when they became extinct following the arrival of humans.

<span class="mw-page-title-main">Dwarf elephant</span> Prehistoric elephant species

Dwarf elephants are prehistoric members of the order Proboscidea which, through the process of allopatric speciation on islands, evolved much smaller body sizes in comparison with their immediate ancestors. Dwarf elephants are an example of insular dwarfism, the phenomenon whereby large terrestrial vertebrates that colonize islands evolve dwarf forms, a phenomenon attributed to adaptation to resource-poor environments and lack of predation and competition.

<i>Palaeoloxodon recki</i> Extinct species of elephant

Palaeoloxodon recki, often known by the synonym Elephas recki is an extinct species of elephant native to Africa and West Asia from the Pliocene or Early Pleistocene to the Middle Pleistocene. During most of its existence, the species represented the dominant elephant species in East Africa. The species is divided into five roughly chronologically successive subspecies. While the type and latest subspecies P. recki recki as well as the preceding P. recki ileretensis are widely accepted to be closely related to Eurasian Palaeoloxodon, the relationships of the other, chronologically earlier subspecies to P. recki recki and P. recki ileretensis are uncertain, with it being suggested they are unrelated and should be elevated to separate species.

<i>Anancus</i> Genus of proboscideans

Anancus is an extinct genus of "tetralophodont gomphothere" native to Afro-Eurasia, that lived from the Tortonian stage of the late Miocene until its extinction during the Early Pleistocene, roughly from 8.5–2 million years ago.

<i>Sinomastodon</i> Extinct genus of gomphothere proboscidean

Sinomastodon is an extinct gomphothere genus known from the Late Miocene to Early Pleistocene of Asia, including China, Japan, Thailand, Myanmar, Indonesia and probably Kashmir.

<i>Palaeoloxodon namadicus</i> Extinct species of elephant

Palaeoloxodon namadicus is an extinct species of prehistoric elephant known from the early Middle to Late Pleistocene of the Indian subcontinent, and possibly also elsewhere in Asia. The species grew larger than any living elephant, and some authors have suggested it to have been the largest known land mammal based on extrapolation from fragmentary remains, though these estimates are speculative.

<i>Stegolophodon</i> Genus of extinct elephant-like mammals

Stegolophodon is an extinct genus of stegodontid proboscideans. It lived during the Miocene epoch in Asia. The earliest fossils are known from the Early Miocene, with one of the oldest fossils being from Japan, estimated to be 17.3 million years old. It is suggested to be the ancestor of Stegodon, and transitional fossils between the two genera known from the Late Miocene of Southeast Asia and Yunnan in South China. Like modern elephants, Stegolophodon developed proal jaw movement, where the lower jaw moves in a back-to-front motion, rather than the oblique chewing motion used by earlier proboscideans, with this development already present by 17.3 million years ago. Members of the genus generally have tetralophodont molars, and retain tusks on the lower jaw. The upper tusks have an enamel band.

<i>Elephas hysudricus</i> Extinct species of mammal

Elephas hysudricus is an extinct elephant species known from the Pleistocene of Asia. It is thought to be ancestral to the living Asian elephant, from which it is distinguished by the molar teeth having a lower crown height and a lower lamellae number. Remains of the species are primarily known from the Indian subcontinent, with the most important remains coming from the Siwalik Hills. The oldest remains of the species in the Siwaliks are placed at around 2.6 million year ago at the beginning of the Early Pleistocene, with the youngest dates in the Siwaliks during the Middle Pleistocene around 0.6 million years ago, though it likely persisted on the subcontinent later than this based on remains found elsewhere. Remains likely attributable to the species are also known from the Levant in Israel and Jordan, dating to the late Middle Pleistocene, likely sometime between 500-100,000 years ago. Isotopic analysis of specimens from the Indian subcontinent suggests that early members of the species were likely primarily grazers, but shifted towards mixed feeding after the arrival of the substantially larger elephant species Palaeoloxodon namadicus to the region. It is suggested to be closely related and possibly ancestral to the extinct Elephas hysudrindicus from the Pleistocene of Java in Indonesia.

<span class="mw-page-title-main">Stegodontidae</span> Family of extinct elephant-like mammals

Stegodontidae is an extinct family of proboscideans from Africa and Asia from the Early Miocene to the Late Pleistocene. It contains two genera, the earlier Stegolophodon, known from the Miocene of Asia and the later Stegodon, from the Late Miocene to Late Pleistocene of Africa and Asia which is thought to have evolved from the former. The group is noted for their plate-like lophs on their teeth, which are similar to elephants and different from those of other extinct proboscideans like gomphotheres and mammutids, with both groups having a proal jaw movement utilizing forward strokes of the lower jaw. These similarities with modern elephants were probably convergently evolved. Like elephantids, stegodontids are thought to have evolved from gomphothere ancestors.

<i>Palaeoloxodon naumanni</i> Extinct species of elephant

Palaeoloxodon naumanni, occasionally called Naumann's elephant, is an extinct species belonging to the genus Palaeoloxodon found in the Japanese archipelago during the Middle to Late Pleistocene around 330,000 to 24,000 years ago. It is named after Heinrich Edmund Naumann who discovered the first fossils at Yokosuka, Kanagawa, Japan. Fossils attributed to P. naumanni are also known from China and Korea, though the status of these specimens is unresolved, and some authors regard them as belonging to separate species.

<i>Elephas hysudrindicus</i> Extinct species of mammal

Elephas hysudrindicus, commonly known also as the Blora elephant in Indonesia, is a species of extinct elephant from the Pleistocene of Java. It is anatomically distinct from the Asian elephant, the last remaining species of elephant under the genus Elephas. The species existed from around the end of the Early Pleistocene until the end of the Middle Pleistocene, when it was replaced by the modern Asian elephant, coexisting alongside fellow proboscidean Stegodon trigonocephalus, as well archaic humans belonging to the species Homo erectus.

<i>Stegodon aurorae</i> Species of fossil elephantoid

Stegodon aurorae, also known as the Akebono elephant, is a species of fossil elephantoid known from Early Pleistocene Japan and Taiwan.

<i>Elephas planifrons</i> Extinct species of mammal

Elephas planifrons is an extinct species of elephant, known from the Late Pliocene-Early Pleistocene of the Indian subcontinent.

<i>Stegoloxodon</i> Extinct genus of dwarf elephant

Stegoloxodon is an extinct genus of dwarf elephant known from the Early Pleistocene of Indonesia. It contains two species, S. indonesicus from Java, and S. celebensis from Sulawesi. Its relationship with other elephants is uncertain.

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  22. 1 2 Shoshani, J.; Tassy, P. (2005). "Advances in proboscidean taxonomy & classification, anatomy & physiology, and ecology & behavior". Quaternary International. 126–128: 5–20. Bibcode:2005QuInt.126....5S. doi:10.1016/j.quaint.2004.04.011.
  23. ↑ Wu, Yan; Deng, Tao; Hu, Yaowu; Ma, Jiao; Zhou, Xinying; Mao, Limi; Zhang, Hanwen; Ye, Jie; Wang, Shi-Qi (16 May 2018). "A grazing Gomphotherium in Middle Miocene Central Asia, 10 million years prior to the origin of the Elephantidae". Scientific Reports. 8 (1): 7640. Bibcode:2018NatSR...8.7640W. doi:10.1038/s41598-018-25909-4. ISSN   2045-2322. PMC   5956065 . PMID   29769581.
  24. ↑ Konidaris, George E.; Tsoukala, Evangelia (2022), Vlachos, Evangelos (ed.), "The Fossil Record of the Neogene Proboscidea (Mammalia) in Greece", Fossil Vertebrates of Greece Vol. 1, Cham: Springer International Publishing, pp. 299–344, doi:10.1007/978-3-030-68398-6_12, ISBN   978-3-030-68397-9, S2CID   245023119 , retrieved 28 February 2023
  25. ↑ Lister, Adrian M.; Dirks, Wendy; Assaf, Amnon; Chazan, Michael; Goldberg, Paul; Applbaum, Yaakov H.; Greenbaum, Nathalie; Horwitz, Liora Kolska (September 2013). "New fossil remains of Elephas from the southern Levant: Implications for the evolutionary history of the Asian elephant". Palaeogeography, Palaeoclimatology, Palaeoecology. 386: 119–130. Bibcode:2013PPP...386..119L. doi:10.1016/j.palaeo.2013.05.013.
  26. ↑ Saegusa, Haruo; Thasod, Yupa; Ratanasthien, Benjavun (2005). "Notes on Asian stegodontids". Quaternary International. 126–128: 31–48. Bibcode:2005QuInt.126...31S. doi:10.1016/j.quaint.2004.04.013.
  27. ↑ van den Bergh, Gerrit D.; Alloway, Brent V.; Storey, Michael; Setiawan, Ruly; Yurnaldi, Dida; Kurniawan, Iwan; Moore, Mark W.; Jatmiko; Brumm, Adam; Flude, Stephanie; Sutikna, Thomas; Setiyabudi, Erick; Prasetyo, Unggul W.; Puspaningrum, Mika R.; Yoga, Ifan (October 2022). "An integrative geochronological framework for the Pleistocene So'a basin (Flores, Indonesia), and its implications for faunal turnover and hominin arrival". Quaternary Science Reviews. 294: 107721. Bibcode:2022QSRv..29407721V. doi:10.1016/j.quascirev.2022.107721. hdl: 10072/418777 . S2CID   252290750.
  28. ↑ Jukar, A. M.; Lyons, S. K.; Wagner, P. J.; Uhen, M. D. (15 January 2021). "Late Quaternary extinctions in the Indian Subcontinent". Palaeogeography, Palaeoclimatology, Palaeoecology. 562: 110137. Bibcode:2021PPP...56210137J. doi: 10.1016/j.palaeo.2020.110137 . ISSN   0031-0182.
  29. 1 2 Samuel T. Turvey, Haowen Tong, Anthony J. Stuart and Adrian M. Lister (2013). "Holocene survival of Late Pleistocene megafauna in China: a critical review of the evidence". Quaternary Science Reviews. 76: 156–166. Bibcode:2013QSRv...76..156T. doi:10.1016/j.quascirev.2013.06.030.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  30. ↑ H. Saegusa, "Comparisons of Stegodon and Elephantid Abundances in the Late Pleistocene of Southern China" Archived 2006-05-08 at the Wayback Machine , The World of Elephants – Second International Congress, (Rome, 2001), 345–349.
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