Proboscidea

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Proboscidea
Temporal range: Middle
Ma
Proboscidean diversity:
Loxodonta cyclotis
Skeleton of Moeritherium
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Mirorder: Tethytheria
Order: Proboscidea
Illiger, 1811
Subclades

Proboscidea (

Ancient Greek προβοσκίς (proboskís) 'elephant's trunk') 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.[1] Three species of elephant are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant
.

Extinct members of Proboscidea include the

paraceratheres, the otherwise largest known land mammals, though this estimate was made based on a single fragmentary femur and is speculative.[2] The largest extant proboscidean is the African bush elephant, with a record of size of 4 m (13.1 ft) at the shoulder and 10.4 t (11.5 short tons).[2]
In addition to their enormous size, later proboscideans are distinguished by tusks and long, muscular trunks, which were less developed or absent in early proboscideans.

Evolution

Over 180 extinct members of Proboscidea have been described.[3] The earliest proboscideans, Eritherium and Phosphatherium are known from the late Paleocene of Africa.[4] The Eocene included Numidotherium, Moeritherium and Barytherium from Africa. These animals were relatively small and some, like Moeritherium and Barytherium were probably amphibious.[5][6]

A major event in proboscidean evolution was the collision of Afro-Arabia with Eurasia, during the Early

Mammuthus (mammoths) migrated out of Africa during the late Pliocene, around 3.6 to 3.2 million years ago.[9]

Over the course of the

Great American interchange,[10] and mammoths migrating into North America around 1.5 million years ago.[11] At the end of the Early Pleistocene, around 800,000 years ago the elephantid genus Palaeoloxodon dispersed outside of Africa, becoming widely distributed in Eurasia.[12] By the beginning of the Late Pleistocene, proboscideans were represented by around 23 species. Proboscideans underwent a dramatic decline during the Late Pleistocene as part of the Late Pleistocene megafauna extinctions, with all remaining non-elephantid proboscideans (including Stegodon, mastodons, and the American gomphotheres Cuvieronius and Notiomastodon) and Palaeoloxodon becoming extinct, with mammoths only surviving in relict populations on islands around the Bering Strait into the Holocene, with their latest survival being on Wrangel Island around 4,000 years ago.[7][13]

The following cladogram is based on endocasts[14]

Proboscidea

Phosphatherium esculliei

Numidotherium koholense

Moeritherium lyonsi

Deinotheriidae

Elephantiformes

Palaeomastodon beadnelli

Elephantimorpha
Mammutida
 

Mammut americanum

Zygolophodon borsoni

Choerolophodon pentelici

Gomphotherium augustidens

Cuvieronius andium

Stegomastodon humboldti

Elephantoidea

Stegodon insignis

Elephantidae

Mammuthus meridionalis

Mammuthus primigenius

Mammuthus columbi

Elephas maximus

Loxodonta africana

Palaeoloxodon

Palaeoloxodon antiquus

Palaeoloxodon falconeri

"plesielephantiforms"
"mastodonts"

Morphology

Over the course of their evolution, proboscideans experienced a significant increase in body size. Some members of the families Deinotheriidae, Mammutidae, Stegodontidae and Elephantidae are thought to have exceeded modern elephants in size, with shoulder heights over 4 metres (13 ft) and masses over 10 tonnes (22,000 lb).[15] As with other megaherbivores, including the extinct sauropod dinosaurs, the large size of proboscideans likely developed to allow them to survive on vegetation with low nutritional value.[16] Their limbs grew longer and the feet shorter and broader.[17] The feet were originally plantigrade and developed into a digitigrade stance with cushion pads and the sesamoid bone providing support, with this change developing around the common ancestor of Deinotheriidae and Elephantiformes.[18] Members of Elephantiformes which have retracted nasal regions of the skull indicating the development of a trunk, as well as well-developed tusks on the upper and lower jaws.[19]

The skull grew larger, especially the cranium, while the neck shortened to provide better support for the skull. The increase in size led to the development and elongation of the mobile trunk to provide reach. The number of

dentine. In primitive proboscideans, a band of enamel covers part of the tusk surface, though in many later groups including modern elephants the band is lost, with elephants only having enamel on the tusk tips of juveniles. The upper tusks were initially modest in size, but from the Late Miocene onwards proboscideans developed increasingly large tusks, with the longest ever recorded tusk being 5.02 metres (16.5 ft) long belonging to the mammutid "Mammut" borsoni found in Greece, with some mammoth tusks likely weighing over 200 kilograms (440 lb). The lower tusks are generally smaller than the upper tusks, but could grow to large sizes in some species, like in Deinotherium (which lacks upper tusks), where they could grow over 1.5 metres (4.9 ft) long, the amebelodontid Konobelodon has lower tusks 1.61 metres (5.3 ft) long, with the longest lower tusks ever recorded being from the primitive elephantid Stegotetrabelodon which are around 2.2 metres (7.2 ft) long.[22]

The molar teeth changed from being replaced vertically as in other mammals to being replaced horizontally in the clade Elephantimorpha.[23] While early Elephantimorpha generally had lower jaws with an elongated mandibular symphysis at the front of the jaw with well developed lower tusks/incisors, from the Late Miocene onwards, many groups convergently developed brevirostrine (shortened) lower jaws with vestigial or no lower tusks.[24][25] Elephantids are distinguished from other proboscideans by a major shift in the molar morphology to parallel lophs rather than the cusps of earlier proboscideans, allowing them to become higher crowned (hypsodont) and more efficient in consuming grass.[26]

Dwarfism

Skeleton of Palaeoloxodon falconeri, one of the smallest known dwarf elephants, with an adult shoulder height less than 1 metre (3.3 ft)

Several species of proboscideans lived on islands and experienced

Channel Islands of California, and several islands of the Mediterranean.[27]

Channel Islands and evolved into the pygmy mammoth. This species reached a height of 1.2–1.8 m (4–6 ft) and weighed 200–2,000 kg (440–4,410 lb). A population of small woolly mammoths survived on Wrangel Island as recently as 4,000 years ago.[27] After their discovery in 1993, they were considered dwarf mammoths.[28] This classification has been re-evaluated and since the Second International Mammoth Conference in 1999, these animals are no longer considered to be true "dwarf mammoths".[29]

Ecology

It has been suggested that members of Elephantimorpha, including mammutids,[30] gomphotheres,[31] and stegodontids,[32] lived in herds like modern elephants. Analysis of remains of the American mastodon (Mammut americanum) suggest that like modern elephants, that herds consisted of females and juveniles and that adult males lived solitarily or in small groups, and that adult males periodically engaged in fights with other males during periods similar to musth found in living elephants. These traits are suggested to be inherited from the last common ancestor of elephantimorphs,[30] with musth-like behaviour also suggested to have occurred in gomphotheres.[33] All elephantimorphs are suggested to have been capable of communication via infrasound, as found in living elephants.[34]

Classification

Below is an unranked taxonomy of proboscidean genera as of 2019.[35][36][37][38]

References

  1. ^ Illiger, Johann Karl Wilhelm (1811). Prodromus Systematis Mammalium et Avium: Additis Terminis Zoographicis Utriusque Classis, Eorumque Versione Germanica. Berolini: Sumptibus C. Salfeld. p. 62.
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  8. ^ H. Saegusa, H. Nakaya, Y. Kunimatsu, M. Nakatsukasa, H. Tsujikawa, Y. Sawada, M. Saneyoshi, T. Sakai Earliest elephantid remains from the late Miocene locality, Nakali, Kenya Scientific Annals, School of Geology, Aristotle University of Thessaloniki, Greece VIth International Conference on Mammoths and Their Relatives, vol. 102, Grevena -Siatista, special volume (2014), p. 175
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  14. , retrieved 22 May 2023
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  16. ^ Carpenter, K. (2006). "Biggest of the big: a critical re-evaluation of the mega-sauropod Amphicoelias fragillimus Cope, 1878". In Foster, J.R.; Lucas, S.G. (eds.). Paleontology and Geology of the Upper Jurassic Morrison Formation. New Mexico Museum of Natural History and Science Bulletin. Vol. 36. New Mexico Museum of Natural History and Science. pp. 131–138.
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  18. from the original on 21 March 2023. Retrieved 3 January 2023.
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  27. ^ a b c Sukumar, pp. 31–33.
  28. S2CID 4249191.{{cite journal}}: CS1 maint: multiple names: authors list (link
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  29. ISSN 0923-9308.{{cite journal}}: CS1 maint: multiple names: authors list (link
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  34. , retrieved 20 April 2024
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Bibliography