Mammuthus meridionalis Temporal range: Early Pleistocene | |
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Mounted skeleton of the Durfort mammoth , National Museum of Natural History, France (Muséum national d'histoire naturelle) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Proboscidea |
Family: | Elephantidae |
Genus: | † Mammuthus |
Species: | †M. meridionalis |
Binomial name | |
†Mammuthus meridionalis (Nesti, 1825) | |
Synonyms | |
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Mammuthus meridionalis, sometimes called the southern mammoth, is an extinct species of mammoth native to Eurasia, including Europe, during the Early Pleistocene, living from around 2.5 million years ago to 800,000 years ago.
Mammuthus meridionalis was originally named by Filippo Nesti in 1825 as Elephas meridionalis based on remains collected from the Upper Valdarno region in Tuscany, Italy. [1]
The taxonomy of extinct elephants was complicated by the early 20th century, and in 1942, Henry Fairfield Osborn's posthumous monograph on the Proboscidea was published, wherein he used various taxon names that had previously been proposed for mammoth species, including replacing Mammuthus with Mammonteus, as he believed the former name to be invalidly published. [2] Mammoth taxonomy was simplified by various researchers from the 1970s onwards, all species were retained in the genus Mammuthus, and many proposed differences between species were instead interpreted as intraspecific variation. [3] The name Archidiskodon meridionalis is retained by some Russian researchers. [4] [5]
M. meridionalis was a large proboscidean, exceeding modern elephants in size. [6] A mature adult male known from a mostly complete skeleton displayed at Forte Spagnolo, L'Aquila, Italy, estimated to be approximately 3.97–4.05 m (13.0–13.3 ft) tall at the shoulder in the flesh, was volumetrically estimated to weigh 10.7–11.4 tonnes (11.8–12.6 short tons). [6] [7] Such sizes are suggested to have been typical for males of this species. [6] Like modern elephants females were considerably smaller, with estimated average adult shoulder height of 3.3 m (10.8 ft) and a weight of around 7 tonnes (7.7 short tons). [8]
The skull was prominently domed, though the height of the dome was lower than later mammoth species. The head represented the highest point of the animal. The body was broad and the back was noticeably sloped. It had robust, elongated twisted tusks, common of mammoths. [8] Its molars had low crowns [9] and around 13 thick enamel ridges (lamellae) on the third molars, substantially lower than the number in later mammoth species. [10] M. meridionalis lived in relatively warm climates, which makes it more probable that it lacked dense fur. [9] The ears are also suggested to have been medium-large sized, with the tail being shorter than living elephants but longer than later mammoth species. [8]
Later European M. meridionalis populations differ from early representatives of the species by having shorter and taller skulls and mandibles, differing shapes of the temporal fossa, orbits and tusk alveoli (sockets), and an increase in the number of lamellae on the teeth and tooth crown height (hypsodonty). [11]
Fossilized plants found with the remains show that M. meridionalis was living in a time of mild climate, generally as warm or slightly warmer than Europe experiences today. Some populations inhabited woodlands, which included oak, ash, beech and other familiar European trees, as well as some that are now exotic to the region, such as hemlock, wing nut and hickory. Further east, discoveries at Ubeidiya (Israel) and Dmanisi (Georgia) show the early mammoth living in a partially open habitat with grassy areas. [9]
Dental microwear of the teeth of M. meridionalis suggest that the species was a variable mixed feeder, that consumed both grass and browse, with its diet varying according to local conditions, [12] with some populations exhibiting browse-dominated feeding, [13] while others grass-dominant. [12]
During the early part of its existence in Europe, it existed alongside the "tetralophodont gomphothere" Anancus arvernensis . Dietary analysis based on microwear suggests that there was niche partitioning between the two species, with M. meridonalis occupying more open habitats. [14]
Juvenile Mammuthus meridionalis have been suggested to have at least occasionally been preyed upon by the large sabertooth cat Homotherium latidens, based on isotopic analysis of specimens from the Venta Micena locality in southeast Spain. [15] Remains from the Fuente Nueva-3 site also in southeast Spain suggests that carcasses of Mammuthus meridionalis were at times scavenged on by the giant hyena Pachycrocuta . [16]
Since many remains of each species of mammoth are known from several localities, it is possible to reconstruct the evolutionary history of the genus through morphological studies. Mammoth species can be identified from the number of enamel ridges (or lamellar plates) on their molars: primitive species had few ridges, and the number increased gradually as new species evolved to feed on more abrasive food items. The crowns of the teeth became deeper in height and the skulls became taller to accommodate this. At the same time, the skulls became shorter from front to back to minimise the weight of the head. [17] [18]
Mammuthus meridionalis is thought to descend from Mammuthus rumanus , the oldest mammoth species known outside of Africa, with the earliest records of M. meridionalis dating to around 2.6-2.5 million years ago, at the beginning of the Pleistocene. [17] Some early members of M. meridionalis spanning from 2.6-2.0 million years ago were historically assigned to the species M. gromovi, which some authors have regarded as the subspecies M. meridionalis gromovi. [17] [11] A population of M. meridionalis evolved into the steppe mammoth (M. trogontherii) with 18–20 third molar ridges in eastern Asia, prior to 1.7 million years ago. [10] The Columbian mammoth (M. columbi) evolved from a population of M. trogontherii that had crossed the Bering Strait and entered North America about 1.5 million years ago, and not M. meridionalis as has been historically suggested. [10] [19] [20] European M. meridionalis specimens from around 2-1.7 million years ago are assigned to the subspecies M. meridionalis meridionalis. Advanced late Early Pleistocene populations of M. meridionalis in Europe, spanning from around 1.7-0.8 million years ago are assigned to the subspecies M. meridionalis vestinus (including the likely synonym M. meridionalis depereti) and M. meridionalistamanensis. These two subspecies may be synonymous with each other. [11] Steppe mammoths replaced M. meridionalis in Europe in a diachronous mosaic pattern at the end of the Early Pleistocene, between around 1 and 0.8-0.7 million years ago, which was also co-incident with the arrival of the straight-tusked elephant (Palaeoloxodon antiquus) into Europe, which may have out-competed M. meridionalis. [17] [11] During the interval of replacement, M. meridionalis and M. trogontherii may have co-existed in some localities, with rare specimens with molar morphology intermediate between the two species suggesting that there may have been hybridisation between them. [21]
The dwarf mammoth species Mammuthus creticus , which inhabited the island of Crete at some point during the Early Pleistocene to early Middle Pleistocene, is suggested to have descended from M. meridionalis. [22]
Remains of M. meridionalis at several sites have been found with cut marks and/or associated with stone tools, suggested to represent evidence of butchery by archaic humans. [23] A number of bones of Mammuthus meridionalis from the Dmanisi site in Georgia, dating to 1.8 million years ago have cut marks likely created by local Homo erectus. [24] At the Fuente Nueva-3 and Barranc de la Boella sites in Spain, dating to approximately 1.3 and 1-0.8 million years ago respectively, remains of M. meridionalis are associated with stone tools (in the latter site of the Acheulean type), primarily lithic flakes. At Barranc de la Boella, some rib bones possibly bear cut marks, [23] with cut marks being definitvely reported from bones found at Fuente Nueva-3. [25] These sites likely represent evidence of opportunistic scavenging, rather than active hunting. [26]
A mammoth is any species of the extinct elephantid genus Mammuthus. They lived from the late Miocene epoch into the Holocene until about 4,000 years ago, with mammoth species at various times inhabiting Africa, Asia, Europe, and North America. Mammoths are distinguished from living elephants by their spirally twisted tusks and in at least some later species, the development of numerous adaptions to living in cold environments, including a thick layer of fur.
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.
A mastodon is a member of the genus Mammut, which was endemic to North America and lived from the late Miocene to the early Holocene. Mastodons belong to the order Proboscidea, the same order as elephants and mammoths. Mammut is the type genus of the extinct family Mammutidae, which diverged from the ancestors of modern elephants at least 27–25 million years ago, during the Oligocene.
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.
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.
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 the largest known species of elephants, over 4 metres (13 ft) tall at the shoulders and over 13 tonnes (29,000 lb) in weight, representing among the largest land mammals ever, 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 like Palaeoloxodon falconeri less than 1 metre (3.3 ft) in shoulder height as fully grown adults, 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.
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.
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.
The Columbian mammoth is an extinct species of mammoth that inhabited North America from southern Canada to Costa Rica during the Pleistocene epoch. The Columbian mammoth descended from Eurasian steppe mammoths that colonised North America during the Early Pleistocene around 1.5–1.3 million years ago, and later experienced hybridisation with the woolly mammoth lineage. The Columbian mammoth was among the last mammoth species, and the pygmy mammoths evolved from them on the Channel Islands of California. The closest extant relative of the Columbian and other mammoths is the Asian elephant.
The Chibanian, more widely known as Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The Chibanian name was officially ratified in January 2020. It is currently estimated to span the time between 0.770 Ma and 0.129 Ma, also expressed as 770–126 ka. It includes the transition in palaeoanthropology from the Lower to the Middle Paleolithic over 300 ka.
The straight-tusked elephant is an extinct species of elephant that inhabited Europe and Western Asia during the Middle and Late Pleistocene. One of the largest known elephant species, mature fully grown bulls on average had a shoulder height of 4 metres (13 ft) and a weight of 13 tonnes (29,000 lb). Straight-tusked elephants likely lived very similarly to modern elephants, with herds of adult females and juveniles and solitary adult males. The species was primarily associated with temperate and Mediterranean woodland and forest habitats, flourishing during interglacial periods, when its range would extend across Europe as far north as Great Britain and eastwards into Russia, while persisting in southern Europe during glacial periods. Skeletons found in association with stone tools and wooden spears suggest they were scavenged and hunted by early humans, including Homo heidelbergensis and their Neanderthal successors.
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.
Mammuthus trogontherii, sometimes called the steppe mammoth, is an extinct species of mammoth that ranged over most of northern Eurasia during the Early and Middle Pleistocene, approximately 1.7 million to 200,000 years ago. The evolution of the steppe mammoth marked the initial adaptation of the mammoth lineage towards cold environments, with the species probably being covered in a layer of fur. One of the largest mammoth species, it evolved in East Asia during the Early Pleistocene, around 1.8 million years ago, before migrating into North America around 1.3 million years ago, and into Europe during the Early/Middle Pleistocene transition, around 1 to 0.7 million years ago. It was the ancestor of the woolly mammoth and Columbian mammoth of the later Pleistocene.
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.
Mammuthus lamarmorai is a species of dwarf mammoth which lived during the late Middle and Late Pleistocene on the island of Sardinia in the Mediterranean. It has been estimated to have had a shoulder height of around 1.4 metres (4.6 ft). Remains have been found across the western part of the island.
Mammuthus africanavus is a species of mammoth known from remains spanning the Late Pliocene-Early Pleistocene found in Central and North Africa in the countries of Chad, Morocco, Tunisia and Algeria. It was originally described by Camille Arambourg in 1952 based on remains found around Lake Ichkeul in north Tunisia as a species of Elephas. Some specimens from this sample may genuinely represent Elephas rather than Mammuthus, though the holotype has been argued to likely represent a true mammoth. Some authors have argued that the species should be placed in Loxodonta, reflecting the difficulty in distinguishing the teeth of early elephantids. It is distinguished from the earlier Mammuthus subplanifrons by having a higher number of ridges/lamellae on the teeth, which display a greater parallelity, the molars being more hypsodont, with the molars having a greater amount of cementum and thinner enamel, and the molar plates exhibit closer spacing.
Mammuthus rumanus is a species of mammoth that lived during the Pliocene in Eurasia. It the oldest mammoth species known outside of Africa.
Villafranchian age is a period of geologic time spanning the Late Pliocene and Early Pleistocene used more specifically with European Land Mammal Ages. Named by Italian geologist Lorenzo Pareto for a sequence of terrestrial sediments studied near Villafranca d'Asti, a town near Turin, it succeeds the Ruscinian age, and is followed by the Galerian.
"Mammut" borsoni is an extinct species of mammutid proboscidean known from the Late Miocene to Early Pleistocene of Eurasia, spanning from western Europe to China. It is the last known mammutid in Eurasia, and amongst the largest of all proboscideans and largest known land mammals.
Phanagoroloxodon is a genus of extinct elephant. It is known from one species, Phanagoroloxodon mammontoides, which is described from a partial skull from Russia, of probable Late Pliocene-Early Pleistocene age.
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