Mammuthus Meridionalis
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''Mammuthus meridionalis'', sometimes called the southern mammoth, is an extinct species of
mammoth A mammoth is any species of the extinct elephantid genus ''Mammuthus.'' They lived from the late Miocene epoch (from around 6.2 million years ago) into the Holocene until about 4,000 years ago, with mammoth species at various times inhabi ...
native to Eurasia during the
Early Pleistocene The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
. Reaching a size exceeding modern elephants, unlike later Eurasian mammoth species, it was largely native to temperate climates and probably did not have a thick layer of fur. First appearing around 2.5 million years ago, probably descending from '' Mammuthus rumanus'', it survived latest in Europe until around 800,000 years ago when it was replaced by the steppe mammoth (''Mammuthus trogontherii'') and the
straight-tusked elephant The straight-tusked elephant (''Palaeoloxodon antiquus'') is an extinct species of elephant that inhabited Europe and Western Asia during the Middle Pleistocene, Middle and Late Pleistocene. One of the largest known elephant species, mature full ...
(''Palaeoloxodon antiquus''). It is the likely ancestor of the dwarf '' Mammuthus creticus'' on the island of Crete.


Taxonomy

''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. The taxonomy of extinct elephants was complicated by the early 20th century, and in 1942, Henry Fairfield Osborn's posthumous monograph on the
Proboscidea 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 l ...
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. 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. The name ''Archidiskodon meridionalis'' is retained by some Russian researchers for the species.


Description

''M. meridionalis'' was a large proboscidean, exceeding modern elephants in size. A mature adult male known from a mostly complete skeleton displayed at Forte Spagnolo, L'Aquila, Italy, estimated to be approximately tall at the shoulder in the flesh, was volumetrically estimated to weigh . Such sizes are suggested to have been typical for males of this species. Like modern elephants females were considerably smaller, with estimated average adult shoulder height of and a weight of around . 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. Its molars had low
crown A crown is a traditional form of head adornment, or hat, worn by monarchs as a symbol of their power and dignity. A crown is often, by extension, a symbol of the monarch's government or items endorsed by it. The word itself is used, parti ...
s and around 13 thick enamel ridges (lamellae) on the third molars, substantially lower than the number in later mammoth species. ''M. meridionalis'' lived in relatively warm climates, which makes it more probable that it lacked dense fur. 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. 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 The temporal fossa is a fossa (shallow depression) on the side of the skull bounded by the temporal lines above, and the zygomatic arch below. Its floor is formed by the outer surfaces of four bones of the skull. The fossa is filled by the te ...
,
orbits In celestial mechanics, an orbit (also known as orbital revolution) is the curved trajectory of an physical body, object such as the trajectory of a planet around a star, or of a natural satellite around a planet, or of an satellite, artificia ...
and tusk alveoli (sockets), and an increase in the number of lamellae on the teeth and tooth crown height (
hypsodont Hypsodont is a pattern of dentition characterized by with high crowns, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on gritty, fibrous material. The oppos ...
y).


Ecology

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 An oak is a hardwood tree or shrub in the genus ''Quercus'' of the beech family. They have spirally arranged leaves, often with lobed edges, and a nut called an acorn, borne within a cup. The genus is widely distributed in the Northern Hemisp ...
, ash,
beech Beech (genus ''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to subtropical (accessory forest element) and temperate (as dominant element of Mesophyte, mesophytic forests) Eurasia and North America. There are 14 accepted ...
and other familiar European trees, as well as some that are now exotic to the region, such as hemlock, wing nut and
hickory Hickory is a common name for trees composing the genus ''Carya'', which includes 19 species accepted by ''Plants of the World Online''. Seven species are native to southeast Asia in China, Indochina, and northeastern India (Assam), and twelve ...
. Further east, discoveries at Ubeidiya (
Israel Israel, officially the State of Israel, is a country in West Asia. It Borders of Israel, shares borders with Lebanon to the north, Syria to the north-east, Jordan to the east, Egypt to the south-west, and the Mediterranean Sea to the west. Isr ...
) and
Dmanisi Dmanisi ( ka, დმანისი, tr, , ) is a town and archaeological site in the Kvemo Kartli region of Georgia (country), Georgia approximately 93 km southwest of the nation’s capital Tbilisi in the river valley of Mashavera. Abandoned i ...
(
Georgia Georgia most commonly refers to: * Georgia (country), a country in the South Caucasus * Georgia (U.S. state), a state in the southeastern United States Georgia may also refer to: People and fictional characters * Georgia (name), a list of pe ...
) show the early mammoth living in a partially open habitat with grassy areas.
Dental microwear Dental microwear analysis is a method to infer diet and behavior in extinct animals, especially in fossil specimens. It has been used on a variety of taxa, including hominids, victoriapithecids, amphicyonids, canids, ursids, hyaenids, hyaenodont ...
of the teeth of ''M. meridionalis'' suggest that the species was a variable mixed feeder, that consumed both grass and
browse Browsing is a kind of orienting strategy. It is supposed to identify something of relevance for the browsing organism. In context of humans, it is a metaphor taken from the animal kingdom. It is used, for example, about people browsing open sh ...
, with its diet varying according to local conditions, with some populations exhibiting browse-dominated feeding, while others grass-dominant. During the early part of its existence in Europe, it existed alongside the "tetralophodont
gomphothere 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 a ...
" '' Anancus arvernensis''. Dietary analysis based on microwear suggests that there was
niche partitioning In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for e ...
between the two species, with ''M. meridonalis'' occupying more open habitats. Other large animals that existed alongside ''Mammuthus meridionalis'' in Early Pleistocene Europe include rhinoceroses of the genus ''
Stephanorhinus ''Stephanorhinus'' is an extinct genus of two-horned rhinoceros native to Eurasia and North Africa that lived during the Late Pliocene to Late Pleistocene. Species of ''Stephanorhinus'' were the predominant and often only species of rhinoceros in ...
,'' the large hippo ''
Hippopotamus antiquus ''Hippopotamus antiquus'' is an extinct species of the genus ''Hippopotamus'' that ranged across Europe during the Early and Middle Pleistocene. It was considerably larger than the living hippopotamus (''Hippopotamus amphibius''). Description ...
,'' giant deer of the genera '' Praemegaceros'' and ''
Eucladoceros ''Eucladoceros'' (Greek language, Greek for "well-branched antler") is an extinct genus of large deer whose fossils have been discovered across Eurasia, from Europe to China, spanning from the Late Pliocene-Early Pleistocene. It is noted for i ...
'', bovines of the genus '' Leptobos'', replaced towards the end of the Early Pleistocene by ''
Bison A bison (: bison) is a large bovine in the genus ''Bison'' (from Greek, meaning 'wild ox') within the tribe Bovini. Two extant taxon, extant and numerous extinction, extinct species are recognised. Of the two surviving species, the American ...
,'' and equines such as '' Equus altidens'' and '' Equus suessenbornensis''. 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. 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 ''Pachycrocuta'' is an extinct genus of hyena. The largest and most well-researched species is ''Pachycrocuta brevirostris'', colloquially known as the giant short-faced hyena as it stood about at the shoulder and it is estimated to have average ...
''.


Evolution

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. ''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 The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
. 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.'' 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. The Columbian mammoth (''M''. ''columbi'') evolved from a population of ''M. trogontherii'' that had crossed the
Bering Strait The Bering Strait ( , ; ) is a strait between the Pacific and Arctic oceans, separating the Chukchi Peninsula of the Russian Far East from the Seward Peninsula of Alaska. The present Russia–United States maritime boundary is at 168° 58' ...
and entered North America about 1.5 million years ago, and not ''M. meridionalis'' as has been historically suggested. 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. meridionalis'' ''tamanensis.'' These two subspecies may be synonymous with each other.'''' 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 The straight-tusked elephant (''Palaeoloxodon antiquus'') is an extinct species of elephant that inhabited Europe and Western Asia during the Middle Pleistocene, Middle and Late Pleistocene. One of the largest known elephant species, mature full ...
(''Palaeoloxodon antiquus'') into Europe, which may have out-competed ''M. meridionalis''.'''' 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. The dwarf mammoth species '' Mammuthus creticus'', which inhabited the island of
Crete Crete ( ; , Modern Greek, Modern: , Ancient Greek, Ancient: ) is the largest and most populous of the Greek islands, the List of islands by area, 88th largest island in the world and the List of islands in the Mediterranean#By area, fifth la ...
at some point during the Early Pleistocene to early Middle Pleistocene, is suggested to have descended from ''M. meridionalis''.''''


Relationship with humans

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 ''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively calle ...
. 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''. 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 Acheulean (; also Acheulian and Mode II), from the French after the type site of Saint-Acheul, is an archaeological industry of stone tool manufacture characterized by the distinctive oval and pear-shaped "hand axes" associated with ''Homo ...
type), primarily
lithic flake In archaeology, a lithic flake is a "portion of rock (geology), rock removed from an objective piece by percussion or pressure,"Andrefsky, W. (2005) ''Lithics: Macroscopic Approaches to Analysis''. 2d Ed. Cambridge, Cambridge University Press and ...
s. At Barranc de la Boella, some rib bones possibly bear cut marks, with cut marks being definitvely reported from bones found at Fuente Nueva-3. These sites likely represent evidence of opportunistic scavenging, rather than active hunting.


References


Bibliography

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External links

{{Taxonbar, from=Q550390 Pleistocene mammals of Asia Mammoths Pleistocene proboscideans Pleistocene mammals of Europe Fossil taxa described in 1825