39-842: Gomphotheres are an extinct group of proboscideans related to modern elephants . First appearing in Africa during the Oligocene , they dispersed into Eurasia and North America during the Miocene and arrived in South America during the Pleistocene as part of the Great American Interchange . Gomphotheres are a paraphyletic group ancestral to Elephantidae , which contains modern elephants, as well as Stegodontidae . While most famous forms such as Gomphotherium had long lower jaws with tusks,
78-540: A paraphyletic Gomphotheriidae. Mammutidae (mastodons) Eritreum Gomphotherium annectens Choerolophodontidae Amebelodontidae (shovel tuskers) Gomphotherium angustidens Gomphotherium steinheimense Elephantoidea ("tetralophodont gomphotheres", Elephantidae) Gomphotherium sylvaticum Gomphotherium inopinatum Gomphotherium browni Gomphotherium tassyi Gomphotherium productum + American gomphotheres Gomphotheres are generally supposed to have been flexible feeders, with
117-460: 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
156-421: A convergent process that occurred multiple times among gomphotheres, as well as other members of Elephantimorpha. The incisors and long lower jaws of primitive gomphotheres were likely used for cutting vegetation, while brevirostrine gomphotheres relied on their trunks to acquire food similar to modern elephants. The limb bones of gomphotheres like those of mammutids are generally more robust than elephantids, with
195-447: A role in their extinction. [REDACTED] Proboscidea Proboscidea ( / ˌ p r oʊ b ə ˈ s ɪ d i ə / ; from Latin proboscis , from 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
234-456: Is suggested to be the ancestor of later New World gomphothere genera. "Trilophodont gomphotheres" dramatically declined during the Late Miocene, likely due to the increasing C 4 grass-dominated habitats, while during the Late Miocene "tetralophodont gomphotheres" were abundant and widespread in Eurasia, where they represented the dominant group of proboscideans. All trilophodont gomphotheres, with
273-829: 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. After their discovery in 1993, they were considered dwarf mammoths. 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". It has been suggested that members of Elephantimorpha, including mammutids, gomphotheres, and stegodontids, lived in herds like modern elephants. Analysis of remains of
312-768: The Channel Islands of California , and several islands of the Mediterranean . 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 . The Columbian mammoth colonised
351-680: The Pleistocene , around or after 2.5 million years ago as part of the Great American Biotic Interchange due to the formation of the Isthmus of Panama , becoming widespread across the continent. The last gomphothere native to Europe, Anancus arvernensis became extinct during the Early Pleistocene, around 1.6–2 million years ago Sinomastodon became extinct at the end of the Early Pleistocene , around 800,000 years ago. From
390-462: The ancestral condition for the group, some later members developed shortened (brevirostrine) lower jaws with either vestigial or no lower tusks and outlasted the long-jawed gomphotheres. This change made them look very similar to modern elephants, an example of parallel evolution . During the Pliocene and Early Pleistocene , the diversity of gomphotheres declined, ultimately becoming extinct outside of
429-524: 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 . Extinct members of Proboscidea include the deinotheres , mastodons , gomphotheres and stegodonts . The family Elephantidae also contains several extinct groups, including mammoths and Palaeoloxodon . Proboscideans include some of
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#1732851503016468-1348: 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. The following cladogram is based on endocasts. Phosphatherium esculliei [REDACTED] Numidotherium koholense [REDACTED] Moeritherium lyonsi [REDACTED] Deinotheriidae [REDACTED] Palaeomastodon beadnelli [REDACTED] Mammut americanum [REDACTED] Zygolophodon borsoni [REDACTED] Choerolophodon pentelici Gomphotherium augustidens [REDACTED] Cuvieronius andium [REDACTED] Stegomastodon humboldti [REDACTED] Stegodon insignis [REDACTED] Mammuthus meridionalis [REDACTED] Mammuthus primigenius [REDACTED] Mammuthus columbi [REDACTED] Elephas maximus [REDACTED] Loxodonta africana [REDACTED] Palaeoloxodon antiquus [REDACTED] Palaeoloxodon falconeri [REDACTED] Over
507-671: 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, with musth-like behaviour also suggested to have occurred in gomphotheres. All elephantimorphs are suggested to have been capable of communication via infrasound , as found in living elephants. Deinotheres may have also lived in herds, based on tracks found in
546-672: The Americas. The last two genera, Cuvieronius ranging southern North America to western South America, and Notiomastodon ranging over most of South America, continued to exist until the end of the Pleistocene around 12,000 years ago, when they became extinct along with many other megafauna species following the arrival of humans . The name "gomphothere" comes from Ancient Greek γόμφος ( gómphos ), "peg, pin; wedge; joint" plus θηρίον ( theríon ), "beast". Gomphotheres differed from elephants in their tooth structure, particularly
585-628: The Bering Land Bridge. Proboscidean groups prominent during the Miocene include the deinotheres , along with the more advanced elephantimorphs , including mammutids (mastodons), gomphotheres , amebelodontids (which includes the "shovel tuskers" like Platybelodon ), choerolophodontids and stegodontids . Around 10 million years ago, the earliest members of the family Elephantidae emerged in Africa, having originated from gomphotheres. The Late Miocene saw major climatic changes, which resulted in
624-510: The Late Miocene of Romania. Over the course of the Neogene and Pleistocene, various members of Elephantida shifted from a browse-dominated diet towards mixed feeding or grazing. Below is a taxonomy of proboscidean genera as of 2019. [REDACTED] Elephantoidea Elephantoidea is a taxonomic group that contains the elephants as well as their closest extinct relatives, including stegodontids and "tetralophodont gomphotheres ",
663-506: The Late Miocene onwards, many groups convergently developed brevirostrine (shortened) lower jaws with vestigial or no lower tusks. 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. Several species of proboscideans lived on islands and experienced insular dwarfism . This occurred primarily during
702-520: 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 ,
741-588: The Pliocene and Pleistocene epochs. In southern North America, Central America and South America, gomphotheres did not become extinct until shortly after the arrival of humans to the Americas, approximately 12,000 years ago, as part of the Late Pleistocene megafauna extinctions of most large mammals across the Americas. Bones of the last gomphothere genera, Cuvieronius and Notiomastodon, dating to shortly before their extinction have been found associated with human artifacts, suggesting that hunting may have played
780-570: The chewing surfaces on the molar teeth. The teeth are considered to be bunodont , that is, having rounded cusps. They are thought to have chewed differently from modern elephants, using an oblique movement (combining back to front and side to side motion) over the teeth rather than the proal movement (a forwards stroke from the back to the front of the lower jaws) used by modern elephants and stegodontids. Like modern elephants and other members of Elephantimorpha , gomphotheres had horizontal tooth replacement, where teeth would progressively migrate towards
819-497: The course of the Early Pleistocene , all non-elephantid probobscideans outside of the Americas became extinct (including mammutids, gomphotheres and deinotheres), with the exception of Stegodon . Gomphotheres dispersed into South America during this era as part of the Great American interchange , and mammoths migrating into North America around 1.5 million years ago. At the end of the Early Pleistocene, around 800,000 years ago
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#1732851503016858-480: 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), with average fully grown males of the mammutid "Mammut" borsoni having an estimated body mass of 16 tonnes (35,000 lb), making it one
897-436: The decline and extinction of many proboscidean groups such as amebelodontids and choerolophodontids. The earliest members of modern genera of Elephantidae appeared during the latest Miocene-early Pliocene around 6-5 million years ago. The elephantid genera Elephas (which includes the living Asian elephant) and Mammuthus (mammoths) migrated out of Africa during the late Pliocene, around 3.6 to 3.2 million years ago. Over
936-480: The elephantid genus Palaeoloxodon dispersed outside of Africa, becoming widely distributed in Eurasia. 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
975-529: The exception of the Asian Sinomastodon , became extinct in Eurasia by the beginning of the Pliocene , along with the global extinction of the "shovel tusker" amebelodontids. The last gomphotheres in Africa, represented by the "tetralophodont gomphothere" genus Anancus , became extinct around the end of the Pliocene and beginning of the Pleistocene. The New World gomphothere genera Notiomastodon and Cuvieronius dispersed into South America during
1014-422: The feet shorter and broader. 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 . Members of Elephantiformes which have retracted nasal regions of the skull indicating the development of a trunk, as well as well-developed tusks on
1053-454: The front of the jaws before they were taken place by more posterior teeth. Unlike modern elephants, many gomphotheres retained permanent premolar teeth though they were absent in some gomphothere genera. Earlier gomphotheres had lower jaws with an elongate mandibular symphysis and lower tusks, the primitive condition for members of Elephantimorpha . Later members developed shortened (brevirostrine) lower jaws and/or vestigial or no lower tusks,
1092-477: The group which contains modern elephants, as well as Stegodontidae . While the North American long jawed proboscideans Gnathabelodon , Eubelodon and Megabelodon been assigned to Gomphotheriidae in some studies other studies suggest that they should be assigned to Amebelodontidae ( Eubelodon, Megabelodon ) or Choerolophodontidae ( Gnathabelodon ). Cladogram of Elephantimorpha after Li et al. 2023, showing
1131-559: The largest and perhaps the largest land mammal ever, with a fragmentary specimen of the Indian elephant species Palaeoloxodon namadicus only known from a partial femur being speculatively estimated in the same study to have possibly reached a body mass of 22 tonnes (49,000 lb). 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. Their limbs grew longer and
1170-807: The largest known land mammals, with the elephant Palaeoloxodon namadicus and mastodon "Mammut" borsoni suggested to have body masses surpassing 16 tonnes (35,000 lb), rivalling or exceeding paraceratheres (the otherwise largest known land mammals) in size. The largest extant proboscidean is the African bush elephant, with a world record of size of 4 m (13.1 ft) at the shoulder and 10.4 t (11.5 short tons). 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. Over 180 extinct members of Proboscidea have been described. The earliest proboscideans, Eritherium and Phosphatherium are known from
1209-586: The late Paleocene of Africa. The Eocene included Numidotherium , Moeritherium and Barytherium from Africa. These animals were relatively small and some, like Moeritherium and Barytherium were probably amphibious. A major event in proboscidean evolution was the collision of Afro-Arabia with Eurasia, during the Early Miocene , around 18-19 million years ago allowing proboscideans to disperse from their African homeland across Eurasia, and later, around 16-15 million years ago into North America across
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1248-491: The latter half of the Early Pleistocene onwards, gomphotheres were extirpated from most of North America, likely due to competition with mammoths and mastodons . The extinction of gomphotheres in Afro-Eurasia has generally been supposed to be the result the expansion of Elephantidae and Stegodon . The morphology of elephantid molars being more efficient than gomphotheres in consuming grass, which became more abundant during
1287-750: The legs also tending to be proportionally shorter. Their bodies also tend to be more proportionally elongate than those of living elephants. "Gomphotheres" are assigned to their own family, Gomphotheriidae, but are widely agreed to be a paraphyletic group. The families Choerolophodontidae and Amebelodontidae (the latter of which includes "shovel tuskers" with flattened lower tusks like Platybelodon ) are sometimes considered gomphotheres sensu lato, though some authors argue that Amebelodontidae should be sunk into Gomphotheriidae. Gomphotheres are divided into two informal groups, "trilophodont gomphotheres", and "tetralophodont gomphotheres". "Tetralophodont gomphotheres" are distinguished from "trilophodont gomphotheres" by
1326-452: The longest lower tusks ever recorded being from the primitive elephantid Stegotetrabelodon which are around 2.2 metres (7.2 ft) long. The molar teeth changed from being replaced vertically as in other mammals to being replaced horizontally in the clade Elephantimorpha . 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
1365-532: 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
1404-778: The mid- Oligocene , with remains from the Shumaysi Formation in Saudi Arabia dating to around 29-28 million years ago. Gomphotheres were uncommon in Afro-Arabia during the Oligocene. Gomphotheres arrived in Eurasia after the connection of Afro-Arabia and Eurasia during the Early Miocene around 19 million years ago, in what is termed the " Proboscidean Datum Event ". Gomphotherium arrived in North America around 16 million years ago, and
1443-428: The presence of four ridges on the fourth premolar and on the first and second molars, rather than the three present in trilophodont gomphotheres. Some authors choose to exclude "tetralophodont gomphotheres" from Gomphotheriidae, and instead assign them to the group Elephantoidea . "Tetralophodont gomphotheres" are thought to have evolved from "trilophodont gomphotheres", and are suggested to be ancestral to Elephantidae ,
1482-605: The upper and lower jaws. 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. In Elephantiformes, the second upper incisor and lower incisor were transformed into ever growing tusks . The tusks are proportionally heavy for their size, being primarily composed of dentine . In primitive proboscideans,
1521-492: The various species having differing browsing , mixed feeding and grazing diets, with the dietary preference of individual species and populations being shaped by local factors such as climatic conditions and competition. Analysis of the tusks of a male Notiomastodon individual suggest that it underwent musth , similar to modern elephants. Notiomastiodon is also suggested to have lived in social family groups, like modern elephants. Gomphotheres originated in Afro-Arabia during
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