Narrow-nosed rhinoceros Temporal range: | |
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Skull of Stephanorhinus hemitoechus | |
Stephanorhinus hemitoechus life restoration | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Perissodactyla |
Family: | Rhinocerotidae |
Genus: | † Stephanorhinus |
Species: | †S. hemitoechus |
Binomial name | |
†Stephanorhinus hemitoechus Falconer, 1859 | |
Range of Stephanorhinus hemitoechus (blue) and Stephanorhinus kirchbergensis (red), with overlapping range in purple | |
Synonyms | |
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The narrow-nosed rhinoceros (Stephanorhinus hemitoechus), also known as the steppe rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus that lived in western Eurasia, including Europe, as well as North Africa [1] during the Pleistocene. It first appeared in Europe around 500,000 years ago during the Middle Pleistocene and survived there until at least 34,000 years Before Present. It was native to temperate and Mediterranean environments, where it fed on low growing plants and to a lesser extent woody plants. Evidence has been found that it was exploited for food by archaic humans, including Neanderthals.
The species was originally described by Hugh Falconer in 1859 based on remains found in cave deposits in Glamorganshire, south Wales in Great Britain, dating to the Eemian/Last interglacial, around 130-115,000 years ago. The species like other Stephanorhinus species was previously assigned historically to the genus Dicerorhinus , which includes the Sumatran rhinoceros, [2] which genetic evidence indicates is the closest living relative of Stephanorhinus. [3]
The narrow-nosed rhinoceros was a large animal, reaching a body mass of approximately 1,500 kilograms (3,300 lb), making it around the same size/slightly larger than S. hundsheimensis, but smaller than S. kirchbergensis (Merck's rhinoceros) and the woolly rhinoceros (Coelodonta antiquitatis). [4] The size of the species was somewhat variable depending on local conditions. [5] [6] The skull is low slung, with the cranium being downwardly directed. [7] The nasal notch (narial incison) is deep, extending back as far as the first upper molar (M1). [8] The teeth are relatively high crowned (hypsodont) compared to other Stephanorhinus species, with the third molars being relatively enlarged. [7] The teeth are relatively shifted forwards in the jaw. [8] Like other species of Stephanorhinus, S. hemitoechus is thought to have borne two horns, [9] though the attachment regions for the horns are "weakly developed" similar to S. etruscus but unlike S. hundsheimensis. [8] The lower jaw is relatively slender in comparison to S. kirchbergensis, and tapers, becoming slender and narrow towards the front (the mandibular symphysis). [8] Compared to the earlier species S. jeanvireti, S. etruscus, and S. hundsheimensis the limb joints are relatively shallow, and the limb bones relatively broad and short. [7]
In comparison to the widespread Merck's rhinoceros (Stephanorhinus kirchbergensis), the narrow-nosed rhinoceros had a less geographically extensive distribution, including much of Europe, [10] [11] as well as West Asia [12] and North Africa. [1] In Europe the species is known from abundant remains in the Iberian Peninsula in the west (where S. kirchbergensis appears to have been rare or absent), [13] eastwards to Italy, [5] Bulgaria [14] and Greece. [15] [16] Its range extended northwards into northern France, [17] Britain (as far north as North Yorkshire [18] ), Germany [2] and Slovakia [19] during warm interglacial periods. [20]
In North Africa, where the species was previously known as Rhinoceros subinermis [15] , remains are known from Cyrenacia in northeast Libya [1] as well as the Maghreb in Morocco [1] and northern Algeria. [21] In West Asia, the range of the species extends from the Levant, including Israel, [22] Palestine, Lebanon, [16] Jordan [23] and Syria [24] in the west, to western Iran, [25] [26] and Azerbaijan in the east, [27] where some remains were previously referred to as the species Rhinoceros binagadensis. [15]
The morphology of the skull suggests that the species was adapted towards a grazing diet. [2] Tooth wear analysis suggests that the narrow-nosed rhinoceros had a variable diet tending towards grazing or mixed feeding, and clearly distinct from the more browsing focused diet inferred for S. kirchbergensis on average. [28] [29] [30] [31] Although the species has been referred to as the "steppe rhinoceros" and presumed to have had a preference for open habitats, the species was ecologically plastic, and occurred in both open grasslands and forested environments. [32] In Europe the species was found in temperate as well as cool climates, and seems to have been intolerant of cold climate conditions like those typically inhabited by the woolly rhinoceros. [19] Its dietary flexibility likely helped it tolerate cool glacial periods. [31] In the Middle East the species inhabited warm, open xeric habitats. [22]
The narrow-nosed rhinoceros probably behaved similarly to living rhinoceroses. A 1993 study suggested that the species may have been sedentary (having a home range and not migratory) and territorial, and probably engaged in ritualized confrontations between males, which may have sometimes broken out into full-on fighting. [9] A rib of a narrow-nosed rhinoceros from Neumark Nord in Germany has been found with a healed fracture, which may have been the result of such a fight. [33]
Across its range, the narrow-nosed rhinoceros lived alongside other megafauna species, including both animals living today, like red deer, fallow deer, roe deer, wild boar, wolves, brown bears, leopards, wild horse and bison, and those that are extinct, like European wild ass, aurochs, cave hyena, Irish elk, cave lions and straight-tusked elephants. [34] S. hemitoechus juveniles represented likely prey items for cave lions and cave hyenas at least on occasion. [31]
Finds at a number of sites suggest that the narrow-nosed rhinoceros was exploited for food by archaic humans. Specimens of S. hemitoechus from the Middle Pleistocene (Marine Isotope Stage 12, 478,000-424,000 years ago) Arago Cave (Caune de l'Arago) site in Southern France shows extensive evidence of butchery (presumably by Tautavel Man, which is found at the same site). The ratios of skeletal elements implies that only the parts of the body with the most meat were carried to the site. The profile of ages of rhino bones in the cave resembles natural mortality curves, suggesting that there was not selective hunting, and the fact that marks of other carnivores are rare implies that the carcasses were acquired by hunting or active scavenging. [35] At the Shishan Marsh site in the Azraq Oasis in northeast Jordan, dating to around 250,000 years ago, stone tools at the site have been found to have protein residue from the butchery of rhinoceroses. As S. hemitoechus is the only rhinoceros species known from the site, it is probable that it was the species butchered. [36]
At the late Middle Pleistocene Gran Dolina site in Spain, a handful of S. cf. hemitoechus bones display cut marks. [37] At Biache-Saint-Vaast in northeast France, dating to MIS 7, around 240,000 years ago, [38] remains of at least 33 individuals of S. hemitoechus were found in association with human artifacts, with a significant proportion displaying cut marks. The mortality profile, which is heavily skewed towards juveniles, with no old adults, may suggest selective hunting of juveniles by Neanderthals. [35] At the collapsed cave of Payre in southeast France, dating to the late Middle Pleistocene, numerous remains of rhinoceroses, primarily S. kirchbergensis along with a smaller amount of S. hemitoechus have been found, which display marks indicative of butchery and are suggested to have been accumulated at the site by Neanderthals. The abundance of teeth found at the site (though other skull material is largely absent) suggests that the Neanderthals may have been using them as tools. Mortality profiles found that young and old individuals were the most abundant at the site. [39] The late Middle Pleistocene sites of Great Yeldham and Grays Thurrock in southern Britain (both probably dating to around 300,000 years ago) where remains of S. hemitoechus have been found have also been suggested as butchery sites. [40]
A skull from Cueva Des-Cubierta in central Spain, dating to the early-mid Late Pleistocene (MIS 4-early MIS 3, ~71-43,000 years ago), exhibits fracturing and cut marks consistent with butchery by Neanderthals. The missing pieces of the skull were not found in the cave, suggesting that it had been butchered off-site. It has been proposed that the skull was kept as a hunting trophy along with the skulls of aurochs and bison. [41] Several other sites in Spain demonstrate the exploitation of S. hemitoechus by Neanderthals during the early-mid Late Pleistocene, including Navalmaíllo Rock Shelter (which represents a single individual brought to a hunting camp and butchered) [42] and Abric Romani (minimum of two individuals). [43] Exploitation of narrow-nosed rhinoceros by Neanderthals was relatively infrequent compared to other types of prey. [42]
The earliest remains of the species in Europe date to the early-mid Middle Pleistocene, around 500,000 years ago, [44] with one of the oldest if not the oldest record in Europe being from Campagna Romana near Rome in Central Italy, dating to Marine Isotope Stage 13. [45] It is suggested to have evolved outside of Western Europe before later migrating into the region. The species may have evolved from the earlier Stephanorhinus etruscus , [27] though other authors suggest it may have evolved from S. hundsheimensis. [45] In North Africa, remains of the narrow-nosed rhinoceros are known dating between 109-53,000 years ago. [1] In Europe, the narrow-nosed rhinoceros survived latest in the southern parts of its range. The last records in Italy date to around 41,000 years ago, [31] while remains dating to 40,000 years ago are known from Bacho Kiro cave in Bulgaria. [14] In the Iberian Peninsula, the latest directly dated records of the species date to approximately 34,000 years ago (Portalón del Tejadilla and Arbreda in Spain) [46] with indirectly dated remains found at Gruta da Figueira Brava in Portugal possibly dating to somewhat later. [31] In the Levant, the species may have survived as recently as 15,500 years ago based on remains found in Hayonim Cave, Israel. [47] [22] Its extinction in southern Europe was suggested in a 2017 study to be due at least in part to climatic change causing habitat fragmentation resulting in population fragmentation, with small populations more likely to become extinct for a variety of reasons, "including loss of genetic variability, inbreeding, genetic drift, demographic fluctuations and environmental variations or natural catastrophes". The study found little evidence of systematic hunting of narrow-nosed rhinoceroses by humans at the time of its extinction. [31]
Elasmotherium is an extinct genus of large rhinoceros that lived in Eastern Europe, Central Asia and East Asia during Late Miocene through to the Late Pleistocene, with the youngest reliable dates around 39,000 years ago. It was the last surviving member of Elasmotheriinae, a distinctive group of rhinoceroses separate from the group that contains living rhinoceros (Rhinocerotinae).
The woolly rhinoceros is an extinct species of rhinoceros that inhabited northern Eurasia during the Pleistocene epoch. The woolly rhinoceros was a member of the Pleistocene megafauna. The woolly rhinoceros was covered with long, thick hair that allowed it to survive in the extremely cold, harsh mammoth steppe. It had a massive hump reaching from its shoulder and fed mainly on herbaceous plants that grew in the steppe. Mummified carcasses preserved in permafrost and many bone remains of woolly rhinoceroses have been found. Images of woolly rhinoceroses are found among cave paintings in Europe and Asia. The range of the woolly rhinoceros contracted towards Siberia beginning around 17,000 years ago, with the youngest known records being around 14,000 years old in northeast Siberia, coinciding with the Bølling–Allerød warming, which likely disrupted its habitat, with environmental DNA records possibly extending the range of the species around 9,800 years ago. Its closest living relative is the Sumatran rhinoceros.
Dicerorhinus is a genus of the family Rhinocerotidae, consisting of a single extant species, the two-horned Sumatran rhinoceros, and several extinct species. The genus likely originated from the Late Miocene of central Myanmar. Many species previously placed in this genus probably belong elsewhere.
Rhinoceros is a genus comprising one-horned rhinoceroses. This scientific name was proposed by Swedish taxonomist Carl Linnaeus in 1758. The genus contains two species, the Indian rhinoceros and the Javan rhinoceros. Although both members are threatened, the Javan rhinoceros is one of the most endangered large mammals in the world with only 60 individuals surviving in Java (Indonesia). The word 'rhinoceros' is of Greek origin meaning "nose-horn".
Stegodon is an extinct genus of proboscidean, related to elephants. It was originally assigned to the family Elephantidae along with modern elephants but is now placed in the extinct family Stegodontidae. Like elephants, Stegodon had teeth with plate-like lophs that are different from those of more primitive proboscideans like gomphotheres and mammutids. Fossils of the genus are known from Africa and across much of Asia, as far southeast as Timor. The oldest fossils of the genus are found in Late Miocene strata in Asia, likely originating from the more archaic Stegolophodon, subsequently migrating into Africa. While the genus became extinct in Africa during the Pliocene, Stegodon persisted in South, Southeast and Eastern Asia into the Late Pleistocene.
Crocuta is a genus of hyena containing the largest extant member of the family, the spotted hyena (Crocuta crocuta). Several fossil species are also known, with the Pleistocene Eurasian cave hyenas either being regarded as distinct species or subspecies of the spotted hyena.
Panthera spelaea, commonly known as the cave lion, is an extinct Panthera species that was native to Eurasia and northwest North America during the Pleistocene epoch. Genetic analysis of ancient DNA has revealed that while closely related, it was a distinct species genetically isolated from the modern lion, with the genetic divergence between the two species estimated at around 500,000 years ago. The earliest fossils of the P. spelaea lineage in Eurasia date to around 700,000 years ago. It is closely related and probably ancestral to the American lion. The species ranged from Western Europe to eastern Beringia in North America, and was a prominent member of the mammoth steppe fauna, and an important apex predator across its range. It became extinct about 13,000 years ago. It closely resembled living lions with a coat of yellowish-grey fur though unlike extant lions, males appear to have lacked manes.
Coelodonta is an extinct genus of Eurasian rhinoceroses that lived from about 3.7 million years to 14,000 years ago, in the Pliocene and the Pleistocene epochs. It is best known from the type species, the woolly rhinoceros, which ranged throughout northern Eurasia during the Pleistocene. The earliest known species, Coelodonta thibetana, lived in Tibet during the Pliocene, with the genus spreading to the rest of Eurasia during the Pleistocene.
The European wild ass or hydruntine is an extinct equine from the Middle Pleistocene to Late Holocene of Europe and West Asia, and possibly North Africa. It is a member of the subgenus Asinus, and closely related to the living Asiatic wild ass. The specific epithet, hydruntinus, means from Otranto.
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 Denmark and eastwards into Russia, while persisting in southern Europe during glacial periods. Skeletons found in association with stone tools and in one case, a wooden spear, suggest they were scavenged and hunted by early humans, including Homo heidelbergensis and their Neanderthal successors.
Panthera fossilis is an extinct species of cat belonging to the genus Panthera, known from remains found in Eurasia spanning the Middle Pleistocene and possibly into the Early Pleistocene.
Cave hyena are extinct species or subspecies of hyena known from Eurasia, which ranged from Western Europe to eastern Asia and Siberia during the Pleistocene epoch. It is well represented in many European caves, primarily dating to the Last Glacial Period. It was an apex predator that preyed on large mammals, and was responsible for the accumulation of hundreds of large Pleistocene mammal bones in areas including horizontal caves, sinkholes, mud pits, and muddy areas along rivers.
Swanscombe Skull Site or Swanscombe Heritage Park is a 3.9-hectare (9.6-acre) geological Site of Special Scientific Interest in Swanscombe, north-west Kent, England. It contains two Geological Conservation Review sites and a National Nature Reserve. The park lies in a former gravel quarry, Barnfield Pit, which is the most important site in the Swanscombe complex, alongside several other nearby pits.
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 much of temperate Eurasia, especially Europe, for most of the Pleistocene. The last two species of Stephanorhinus – Merck's rhinoceros and the narrow-nosed rhinoceros – went extinct during the last glacial period.
Stephanorhinus kirchbergensis, also known as Merck's rhinoceros is an extinct species of rhinoceros belonging to the genus Stephanorhinus from the Early-Middle to Late Pleistocene of Eurasia. Its range spanned from Western Europe to Eastern Asia. Among the last members of the genus, it co-existed alongside Stephanorhinus hemitoechus in the western part of its range.
Bubalus murrensis, also known as European water buffalo, is an extinct water buffalo species native to Europe during the Pleistocene epoch, possibly persisting into the Holocene.
The Clacton Spear, or Clacton Spear Point, is the tip of a wooden spear discovered in Clacton-on-Sea in 1911. At approximately 400,000 years old, it is the oldest known worked wooden implement.
The Cave of the Angel refers to several cave-related structures located in the Aras mountain range near the town of Lucena, Córdoba province in Spain. The site contains lithic material of an Acheulean typology and dates to that from the Middle Pleistocene to the Upper Pleistocene. There is geological, paleontological, and archaeological evidence indicating an intense and long-term occupation of this site. The numerous bone and lithic remains found in this site, as well as the matrix surrounding them, reveal exposure to fire. This, along with the finding of a wide and deep combustion structure in the stratigraphic profile, support the hypothesis that at the time of the occupation of the site by humans there was processing and consumption of big mammals.
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Pliorhinus is an extinct genus of rhinoceros known from the Late Miocene and Pliocene of Eurasia. The type species, Pliorhinus megarhinus, was previously assigned to Dihoplus.