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probóscidea

Proboscidea ( / ˌ p r ɒ b ə ˈ s k ɪ d i ə / ; del latín proboscis , del griego antiguo προβοσκίς ( proboskís )  'trompa de elefante') es un orden taxonómico de mamíferos afrotherianos que contiene una familia viva ( Elephantidae ) y varias familias extintas. Descrito por primera vez por J. Illiger en 1811, abarca a los elefantes y sus parientes cercanos. [1] Actualmente se reconocen tres especies de elefante : el elefante africano de sabana, el elefante africano de selva y el elefante asiático .

Los miembros extintos de Proboscidea incluyen los deinotheres , mastodontes , gomphotheres y estegodontos . La familia Elephantidae también contiene varios grupos extintos, incluidos los mamuts y los Palaeoloxodon . Los proboscidios incluyen algunos de los mamíferos terrestres más grandes conocidos. El mamífero terrestre más grande de todos los tiempos pudo haber sido un proboscídeo; Se ha estimado que el elefante Palaeoloxodon namadicus mide hasta 5,2 m (17,1 pies) en el hombro y puede haber pesado hasta 22 t (24,3 toneladas cortas), superando a los paraceratheres , los mamíferos terrestres más grandes conocidos, aunque esta estimación se hizo basado en un único fémur fragmentario y es especulativo. [2] El proboscidio más grande que existe es el elefante africano de sabana , con un tamaño récord de 4 m (13,1 pies) en el hombro y 10,4 t (11,5 toneladas cortas). [2] Además de su enorme tamaño, los proboscídeos posteriores se distinguen por colmillos y troncos largos y musculosos, que estaban menos desarrollados o ausentes en los primeros proboscídeos.

Evolución

Se han descrito más de 180 miembros extintos de Proboscidea. [3] Los primeros proboscidios, Eritherium y Phosphatherium, se conocen del Paleoceno tardío de África. [4] El Eoceno incluyó Numidotherium , Moeritherium y Barytherium de África. Estos animales eran relativamente pequeños y algunos, como Moeritherium y Barytherium, probablemente eran anfibios. [5] [6]

Un acontecimiento importante en la evolución de los proboscidios fue la colisión de Afro-Arabia con Eurasia, durante el Mioceno temprano , hace unos 18-19 millones de años, lo que permitió a los proboscidios dispersarse desde su tierra natal africana a través de Eurasia, y más tarde, hace unos 16-15 millones de años, hacia América del Norte a través del Puente Terrestre de Bering. Los grupos proboscídeos prominentes durante el Mioceno incluyen los deinotheres , junto con los elefantimorfos más avanzados , incluidos mamutidos (mastodontes), gonfoterios , amebelodontidos (que incluye a los "colmillos de pala" como Platybelodon ), corolofodóntidos y estegodóntidos . [7] Hace unos 10 millones de años, los primeros miembros de la familia Elephantidae surgieron en África, originados a partir de gonfoterios. [8] El Mioceno tardío vio cambios climáticos importantes, que resultaron en el declive y extinción de muchos grupos proboscídeos como los amebelodontidos y los corolofodóntidos. [7] Los primeros miembros de los géneros modernos de Elephantidae aparecieron durante el Mioceno tardío y el Plioceno temprano, hace alrededor de 6 a 5 millones de años. Los géneros de elefantidos Elephas (que incluye al elefante asiático vivo) y Mammuthus (mamuts) emigraron de África a finales del Plioceno, hace entre 3,6 y 3,2 millones de años. [9]

En el transcurso del Pleistoceno temprano , todos los probobscidios no elefantes fuera de América se extinguieron (incluidos los mamutidos, gonfoterios y deinotheres), con la excepción de Stegodon . [7] Los gonfoterios se dispersaron en América del Sur durante esta era como parte del Gran Intercambio Americano , [10] y los mamuts migraron a América del Norte hace alrededor de 1,5 millones de años. [11] A finales del Pleistoceno temprano, hace unos 800.000 años, el género de elefantidos Palaeoloxodon se dispersó fuera de África y se distribuyó ampliamente en Eurasia. [12] A principios del Pleistoceno tardío , los proboscídeos estaban representados por alrededor de 23 especies. Los proboscidios sufrieron un dramático declive durante el Pleistoceno tardío como parte de las extinciones de megafauna del Pleistoceno tardío , extinguiéndose todos los proboscidios no elefántidos restantes (incluidos Stegodon , mastodontes y los gonfoterios americanos Cuvieronius y Notiomastodon ) y Palaeoloxodon , y los mamuts solo sobrevivieron en forma relicta. poblaciones en islas alrededor del Estrecho de Bering hasta el Holoceno, y su última supervivencia se produjo en la isla Wrangel hace unos 4.000 años. [7] [13]

El siguiente cladograma se basa en endocasts [14]

Morfología

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 premolars, incisors and canines decreased. The cheek teeth (molars and premolars) became larger and more specialised.[17] In Elephantiformes, the second upper incisor and lower incisor were transformed into ever growing tusks.[20][21] The tusks are proportionally heavy for their size, being primarily composed 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 insular dwarfism. This occurred primarily during the Pleistocene, when some elephant populations became isolated by fluctuating sea levels, although dwarf elephants did exist earlier in the Pliocene. These elephants likely grew smaller on islands due to a lack of large or viable predator populations and limited resources. By contrast, small mammals such as rodents develop gigantism in these conditions. Dwarf proboscideans are known to have lived in Indonesia, the Channel Islands of California, and several islands of the Mediterranean.[27]

Elephas celebensis of Sulawesi is believed to have descended from Elephas planifrons. Elephas falconeri of Malta and Sicily was only 1 m (3 ft), and had probably evolved from the straight-tusked elephant. Other descendants of the straight-tusked elephant existed in Cyprus. Dwarf elephants of uncertain descent lived in Crete, Cyclades and Dodecanese, while dwarf mammoths are known to have lived in Sardinia.[27] The Columbian mammoth colonised the 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] Deinotheres may have also lived in herds, based on tracks found in the Late Miocene of Romania.[35] Over the course of the Neogene and Pleistocene, various members of Elephantida shifted from a browse-dominated diet towards mixed feeding or grazing.[36]

Classification

Below is an unranked taxonomy of proboscidean genera as of 2019.[37][38][39][40]

References

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Bibliography

Wikispecies has information related to Proboscidea.
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