''Stephanorhinus kirchbergensis'', also known as Merck's rhinoceros or the forest rhinoceros, is an extinct species of rhino known from the Middle to

Late Pleistocene The Late Pleistocene is an unofficial Age (geology), age in the international geologic timescale in chronostratigraphy, also known as Upper Pleistocene from a Stratigraphy, stratigraphic perspective. It is intended to be the fourth division of ...

Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...

. One of the last members of the genus ''

Stephanorhinus ''Stephanorhinus'' is an extinct genus of two-horned rhinoceros native to Eurasia and North Africa that lived during the Pliocene to Late Pleistocene. Species of ''Stephanorhinus'' were the predominant and often only species of rhinoceros in much ...

'', it is considered to be a typical component of the

interglacial An interglacial period (or alternatively interglacial, interglaciation) is a geological interval of warmer global average temperature lasting thousands of years that separates consecutive glacial periods within an ice age. The current Holocene in ...

Palaeoloxodon ''Palaeoloxodon'' is an extinct genus of elephant. The genus originated in Africa during the Pliocene era, and expanded into Eurasia during the Pleistocene era. The genus contains some of the largest known species of elephants, over four metres t ...

'' large faunal assemblage in Europe. Among extant species of rhinoceroses it is most closely related to the

Sumatran rhinoceros The Sumatran rhinoceros (''Dicerorhinus sumatrensis''), also known as the Sumatran rhino, hairy rhinoceros or Asian two-horned rhinoceros, is a rare member of the family Rhinocerotidae and one of five extant species of rhinoceros. It is the o ...

, while the well known

woolly rhinoceros The woolly rhinoceros (''Coelodonta antiquitatis'') is an extinct species of rhinoceros that was common throughout Europe and Asia during the Pleistocene epoch and survived until the end of the last glacial period. The woolly rhinoceros was a me ...

was another close relative. In the western part of its range, it was

sympatric In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sh ...

with ''

Stephanorhinus hemitoechus The narrow-nosed rhinoceros (''Stephanorhinus hemitoechus'') is an extinct species of rhinoceros that lived in western Eurasia and North Africa during the Pleistocene. It first appeared in Europe some 600,000 years ago, and survived there until ...

''.

Etymology and taxonomy

The first part of the genus name is derived from that of King Stephen I of Hungary, and the second part from 'rhinos' (ρινος, meaning "

nose A nose is a protuberance in vertebrates that houses the nostrils, or nares, which receive and expel air for respiration alongside the mouth. Behind the nose are the olfactory mucosa and the sinuses. Behind the nasal cavity, air next passes th ...

"), as with ''Dicerorhinus.'' The species name was given by Georg Friedrich von Jäger in 1839 for

Kirchberg an der Jagst Kirchberg an der Jagst is a town in the district of Schwäbisch Hall, in Baden-Württemberg, Germany. It is located on the river Jagst, 11 km northwest of Crailsheim. The Second World War resistance member Friedrich Gustav Jaeger Fried ...

Baden-Württemberg Baden-Württemberg (; ), commonly shortened to BW or BaWü, is a German state () in Southwest Germany, east of the Rhine, which forms the southern part of Germany's western border with France. With more than 11.07 million inhabitants across a ...

, Germany where the type specimens had been found. It is often known in English (and equivalents in other languages) as Merck's rhinoceros after Carl Heinrich Merck, who gave the initial name to the species in 1784 as ''Rhinoceros incisivus'', that is now considered a ''

nomen oblitum In zoological nomenclature, a ''nomen oblitum'' (plural: ''nomina oblita''; Latin for "forgotten name") is a disused scientific name which has been declared to be obsolete (figuratively 'forgotten') in favour of another 'protected' name. In its p ...

'', and who after a widely used

junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...

of the species, ''Rhinoceros/Dicerorhinus mercki'' (historically several alternate spellings) was named by

Johann Jakob Kaup Johann Jakob von Kaup (10 April 1803 – 4 July 1873) was a German naturalist. A proponent of natural philosophy, he believed in an innate mathematical order in nature and he attempted biological classifications based on the Quinarian system. Kaup ...

in 1841.

Description

Merck's rhinoceros is considered to be large for a rhinoceros, with a particularly large specimen from Poland reaching an estimated height at the

withers The withers is the ridge between the shoulder blades of an animal, typically a quadruped. In many species, it is the tallest point of the body. In horses and dogs, it is the standard place to measure the animal's height. In contrast, cattle ar ...

of 182 centimetres. The bones of the skeleton are robust and massive. The skull of Merck's rhinoceros is elongated, with the septum nasalis ossified only towards its anterior (front) end. The

mandibular symphysis In human anatomy, the facial skeleton of the skull the external surface of the mandible is marked in the median line by a faint ridge, indicating the mandibular symphysis (Latin: ''symphysis menti'') or line of junction where the two lateral halves ...

is relatively long and the mandible has a horizontal high, thick branch.Selected records of Stephanorhinus kirchbergensis (Jäger, 1839 (Mammalia, Rhinocerotidae) in Italy
Emmanuel M.E. BILLIA & Carmelo PETRONIO Bollettino della Società Paleontologica Italiana, 48 (1), 2009, xx-xx. Modena

Dental anatomy

The enamel of the teeth is very thick, and often bright coloured and smooth, with very thin or absent coronal cement. The buccal (cheek-facing) sides of the teeth often have sub-vertical bluish lines. Tooth dimensions are highly variable in comparison to other ''Stephanorhinus'' species. The upper teeth, especially the molars, are much higher towards the buccal side than to the lingual (towards the tongue) side. The ectolophs of the first and second upper molars have shallower folds, especially the fold between the paracone and mesostyle, than those of '' S. hemitoechus,'' resulting in a less pronounced undulation. In comparison to other species of ''Stephanorhinus,'' the premolars of ''S. kirchbergensis'' are mesially (towards the front of the tooth) broad and relatively lingually short. The upper premolar ectoloph folds are shallow, and have narrow anterior valleys. The ectoloph curves strongly mesially and often distally (towards the hind portion of the tooth) towards the inside of the tooth. In both upper molars and premolars, the metalophs and the protolophs are distinctly bulbous. The lower premolars and molars are similar and hard to distinguish.

Origin

The origin of the species is obscure, with various authors suggesting either a European or Asian origin. The earliest definitive records are from

Choukoutien Zhoukoudian Area () is a town and an area located on the east Fangshan District, Beijing, China. It borders Nanjiao and Fozizhuang Townships to its north, Xiangyang, Chengguan and Yingfeng Subdistricts to its east, Shilou and Hangcunhe Towns to ...

Locality 13, in

Fangshan District Fangshan District () is situated in the southwest of Beijing, away from downtown Beijing. It has an area of and a population of 814,367 (2000 Census). The district is divided into 8 subdistricts, 14 towns, and 6 townships. The district administ ...

near Beijing at around the Early-Middle Pleistocene transition, though the older records of ''Stephanorhinus lantianensis'' and ''Stephanorhinus yunchuchenensis'' have been suggested to be synonyms of ''S. kirchbergensis'' by some sources. It appears in Europe during the early Middle Pleistocene between 0.7 and 0.6 million years ago, existing alongside the already present ''S. hundsheimensis''. Mitochondrial and nuclear genomes obtained from a permafrost specimen and a dental

proteome The proteome is the entire set of proteins that is, or can be, expressed by a genome, cell, tissue, or organism at a certain time. It is the set of expressed proteins in a given type of cell or organism, at a given time, under defined conditions. ...

suggest that it is more closely related to the

than the

, but its relationship to other ''

'' species remains unclear.

Range

Its range spans from Europe to East Asia, but appears to be absent from the Iberian Peninsula.Billa, E.M.E. 2011a. Occurrences of Stephanorhinus kirchbergensis (Jäger, 1839) (Mammalia, Rhinocerotidae) in Eurasia - An account. Acta Palaeontologica Romaniae 7: 17-40 It is presumed to have had a preference for closed forest and woodland habitats, as opposed the to open grassland habitats favoured by ''S. hemitoechus''. Its range extended into the

Arctic Circle The Arctic Circle is one of the two polar circles, and the most northerly of the five major circles of latitude as shown on maps of Earth. Its southern equivalent is the Antarctic Circle. The Arctic Circle marks the southernmost latitude at w ...

, with a 70–48 thousand-year-old skull known from arctic

Yakutia Sakha, officially the Republic of Sakha (Yakutia),, is the largest republic of Russia, located in the Russian Far East, along the Arctic Ocean, with a population of roughly 1 million. Sakha comprises half of the area of its governing Far Eas ...

in the

Chondon River The Chondon ( rus, Чондон; sah, Чондоон) is a river in Ust-Yansky District, Sakha Republic (Yakutia), Russia. It is long, with a drainage basin of . The Chondon mammoth was discovered in 2013 in the Chondon basin, at the feet of th ...

valley and a late Middle Pleistocene aged lower jaw from the

Yana River The Yana ( rus, Я́на, p=ˈjanə; sah, Дьааҥы, ''Caañı'') is a river in Sakha in Russia, located between the Lena to the west and the Indigirka to the east. Course It is long, and its drainage basin covers . Including its longest ...

valley. Teeth are known from caves in

Primorsky Krai Primorsky Krai (russian: Приморский край, r=Primorsky kray, p=prʲɪˈmorskʲɪj kraj), informally known as Primorye (, ), is a federal subject (a krai) of Russia, located in the Far East region of the country and is a part of the ...

, suggested to date between 50,000 and 25,000 years ago based on dates of other bones found in the deposit, which are the easternmost known records. A tooth of ''S.'' cf''. kirchbergensis'' of an unknown age is known from the

Lut Desert The Lut Desert, widely referred to as Dasht-e Lut ( fa, دشت لوت, "Emptiness Plain"), is a large salt desert located in the provinces of Kerman Province, Kerman and Sistan and Baluchestan Province, Sistan and Baluchestan, Iran. It is the Li ...

in eastern Iran. It is fairly common throughout the Pleistocene in North China, but is a rarer component of South Chinese assemblages, being known from around 30 localities in the region. Antoine (2012) states that ''D.'' ''choukoutienensis'', ''D. lantianensis'', and ''D. yunchuchenensis'' are local names for the taxon, without elaboration. Its range was strongly controlled by glacial cycles, with the species experiencing repeated cycles of expansion and contraction as the ice sheets advanced, this accounts for the relative rarity of its remains in comparison to the woolly rhinoceros. The species' range underwent significant reduction during the Last Glacial Period, with the youngest records from Italy being in

Marine isotope stage Marine isotope stages (MIS), marine oxygen-isotope stages, or oxygen isotope stages (OIS), are alternating warm and cool periods in the Earth's paleoclimate, deduced from oxygen isotope data reflecting changes in temperature derived from data ...

(MIS) 4 and 3. Radiocarbon dated remains from the Altai date to around 40,000 years ago. The youngest reliable records in China are from the Rhino Cave in

Hubei Hubei (; ; alternately Hupeh) is a landlocked province of the People's Republic of China, and is part of the Central China region. The name of the province means "north of the lake", referring to its position north of Dongting Lake. The prov ...

, which is early Late Pleistocene in age. Though less definitive remains are known from near

Harbin Harbin (; mnc, , v=Halbin; ) is a sub-provincial city and the provincial capital and the largest city of Heilongjiang province, People's Republic of China, as well as the second largest city by urban population after Shenyang and largest ...

Heilongjiang Heilongjiang () formerly romanized as Heilungkiang, is a province in northeast China. The standard one-character abbreviation for the province is (). It was formerly romanized as "Heilungkiang". It is the northernmost and easternmost province ...

, which are thought to be 20 kya in age. Records from Migong Cave just south of the Yangtze River in the Three Gorges area are suggested to date to MIS 2 (29,000-14,000 years ago).

Diet

Merck's rhinoceros has been interpreted as a browser, feeding on the branches and leaves of trees and shrubs. It had a more specialised diet than ''S. hundsheimensis'' and was clearly distinct from the grazing diet hypothesised for ''S. hemitoechus''. Despite their morphological differences, dental wear analysis of several European ''S. kirchbergensis'' and ''S. hemitoechus'' populations were similar and indicative of mixed feeding, suggesting dietary convergence due to low habitat variability during the Pleistocene. Analysis of plant material embedded within teeth from the Neumark-Nord locality in Germany found remains of ''

Populus ''Populus'' is a genus of 25–30 species of deciduous flowering plants in the family Salicaceae, native to most of the Northern Hemisphere. English names variously applied to different species include poplar (), aspen, and cottonwood. The we ...

'' (poplar or aspen) ''

Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...

'' (oak), ''

Crataegus ''Crataegus'' (), commonly called hawthorn, quickthorn, thornapple, Voss, E. G. 1985. ''Michigan Flora: A guide to the identification and occurrence of the native and naturalized seed-plants of the state. Part II: Dicots (Saururaceae–Cornacea ...

'' (hawthorn), ''

Pyracantha ''Pyracantha'' (from Greek "fire" and "thorn", hence firethorn) is a genus of large, thorny evergreen shrubs in the family Rosaceae, with common names firethorn or pyracantha. They are native to an area extending from Southwest Europe east to ...

'', ''

Urtica ''Urtica'' is a genus of flowering plants in the family Urticaceae. Many species have stinging hairs and may be called nettles or stinging nettles, although the latter name applies particularly to ''Urtica dioica''. ''Urtica'' species are food f ...

'' (nettles) and ''

Nymphaea ''Nymphaea'' () is a genus of hardy and tender aquatic plants in the family Nymphaeaceae. The genus has a cosmopolitan distribution. Many species are cultivated as ornamental plants, and many cultivars have been bred. Some taxa occur as introduc ...

'' (water lilies) as well as indeterminate remains of

Betulaceae Betulaceae, the birch family, includes six genera of deciduous nut-bearing trees and shrubs, including the birches, alders, hazels, hornbeams, hazel-hornbeam, and hop-hornbeams numbering a total of 167 species. They are mostly natives of the t ...

Rosaceae Rosaceae (), the rose family, is a medium-sized family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus ''Rosa''. Among the most species-rich genera are ''Alchemilla'' (270), ''Sorbus ...

, and

Poaceae Poaceae () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos and the grasses of natural grassland and species cultivated in lawns an ...

(grass). Preserved plant remains found with the teeth on the arctic Chondon skull included twigs of ''Salix'' (

willow Willows, also called sallows and osiers, from the genus ''Salix'', comprise around 400 speciesMabberley, D.J. 1997. The Plant Book, Cambridge University Press #2: Cambridge. of typically deciduous trees and shrubs, found primarily on moist s ...

), ''Betula'' (

birch A birch is a thin-leaved deciduous hardwood tree of the genus ''Betula'' (), in the family Betulaceae, which also includes alders, hazels, and hornbeams. It is closely related to the beech-oak family Fagaceae. The genus ''Betula'' contains 30 ...

) and abundant ''Larix'' (

larch Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains furt ...

) alongside fragments of

Ericaceae The Ericaceae are a family of flowering plants, commonly known as the heath or heather family, found most commonly in acidic and infertile growing conditions. The family is large, with c.4250 known species spread across 124 genera, making it th ...

(heather);

sedges The Cyperaceae are a family of graminoid (grass-like), monocotyledonous flowering plants known as sedges. The family is large, with some 5,500 known species described in about 90 genera, the largest being the "true sedges" genus ''Carex'' wit ...

were notably absent. A specimen from

Eemian The Eemian (also called the last interglacial, Sangamonian, Sangamonian Stage, Ipswichian, Mikulin, Kaydaky, penultimate,NOAA - Penultimate Interglacial Period http://www.ncdc.noaa.gov/global-warming/penultimate-interglacial-period Valdivia or Ri ...

aged deposits in

Gorzów Wielkopolski Gorzów Wielkopolski (; german: Landsberg an der Warthe) often abbreviated to Gorzów Wlkp. or simply Gorzów, is a city in western Poland, on the Warta river. It is the second largest city in the Lubusz Voivodeship with 120,087 inhabitants (Decemb ...

in Poland had twigs of ''Corylus'' (

hazel The hazel (''Corylus'') is a genus of deciduous trees and large shrubs native to the temperate Northern Hemisphere. The genus is usually placed in the birch family Betulaceae,Germplasmgobills Information Network''Corylus''Rushforth, K. (1999). ...

), ''Carpinus'' (

hornbeam Hornbeams are hardwood trees in the flowering plant genus ''Carpinus'' in the birch family Betulaceae. The 30–40 species occur across much of the temperate regions of the Northern Hemisphere. Origin of names The common English name ''hornbeam' ...

), and ''

Viscum ''Viscum'' is a genus of about 70–100 species of mistletoes, native to temperate and tropical regions of Europe, Africa, Asia and Australasia. Traditionally, the genus has been placed in its own family Viscaceae, but recent genetic research by ...

'' (mistletoe)'','' alongside fruit scales of birch, with hazel and birch dominating amongst the pollen. The pollen from a specimen found at

Spinadesco Spinadesco ( Cremunés: ) is a ''comune'' (municipality) in the Province of Cremona in the Italian region Lombardy, located about southeast of Milan and about west of Cremona. Spinadesco borders the following municipalities: Acquanegra Cremonese ...

in Italy was dominated (~50%) by trees, particularly ''Alnus'' (

alder Alders are trees comprising the genus ''Alnus'' in the birch family Betulaceae. The genus comprises about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few sp ...

) and ''Fagus'' (

beech Beech (''Fagus'') is a genus of deciduous trees in the family Fagaceae, native to temperate Europe, Asia, and North America. Recent classifications recognize 10 to 13 species in two distinct subgenera, ''Engleriana'' and ''Fagus''. The ''Engle ...

), with ''

Hippophae rhamnoides ''Hippophae rhamnoides'', also known as sea-buckthorn, is a species of flowering plant in the family Elaeagnaceae, native to the cold-temperate regions of Europe and Asia. It is a spiny deciduous shrub. The plant is used in the food and cosmetic ...

'' (sea buckthorn), dominating amongst the shrubs, with around 30% of the total contribution being from a variety of herbaceous plants.

Human exploitation

At the Taubach

travertine Travertine ( ) is a form of terrestrial limestone deposited around mineral springs, especially hot springs. It often has a fibrous or concentric appearance and exists in white, tan, cream-colored, and even rusty varieties. It is formed by a pro ...

site in

Thuringia Thuringia (; german: Thüringen ), officially the Free State of Thuringia ( ), is a state of central Germany, covering , the sixth smallest of the sixteen German states. It has a population of about 2.1 million. Erfurt is the capital and larg ...

, Germany, which dates to the

(approximately 130,000-115,000 years ago) remains of Merck's rhinoceros with cut marks are known. The vast majority of remains were of young subadults, alongside a much smaller number of adults. It has been suggested that the rhinoceroses were killed and butchered on site by

Neanderthals Neanderthals (, also ''Homo neanderthalensis'' and erroneously ''Homo sapiens neanderthalensis''), also written as Neandertals, are an Extinction, extinct species or subspecies of archaic humans who lived in Eurasia until about 40,000 years ag ...

.Bratlund, B. 1999
Taubach revisited
''Jahrbuch des Römisch-Germanischen Zentralmuseums Mainz'' 46: 61-174.

Gallery

File:Dicerorhinus kirchbergensis.JPG File:Stephanorhinus kirckbergensis skull.jpg, Partial skull in Stuttgart File:Stephanorhinus kirchbergensis.JPG, 300,000 year old dentary fragment from the United Kingdom in the NHM, London File:Naturgeschichte Museum Schaffhausen Waldnashorn aus Flurlingen.jpg, Remains of Merck's rhinoceros from Germany File:Stephanorhinus kirchbergensis skull with horn attachment positions.png, View of the top of the snout of a ''S. kirchbergensis'' skull, showing the rugose texture of the horn attachment area

References

{{Taxonbar, from=Q310972 Pleistocene rhinoceroses Prehistoric mammals of Europe Pleistocene mammals of Asia