Rhinos

from Wikipedia, the free encyclopedia
Rhinos
White rhinos in Namibia

White rhinos in Namibia

Systematics
Class : Mammals (mammalia)
Subclass : Higher mammals (Eutheria)
Superordinate : Laurasiatheria
without rank: Scrotifera
Order : Unpaired ungulate (Perissodactyla)
Family : Rhinos
Scientific name
Rhinocerotidae
Owen , 1845

The rhinoceros (Rhinocerotidae) or rhinoceroses form a family of odd ungulates (Perissodactyla) with five species still alive today. They are characterized by a strong body and short limbs with three toes as well as a large head, which in all representatives living today has a distinctive formation, consisting of one or two horns - which give the family its name. The family is one of the most diverse and successful in the history of mammals and was spread over large parts of Eurasia , Africa and North America during its evolutionary history, which began almost 50 million years ago . Their decline began at the end of the Miocene around 6 to 5 million years ago in connection with climatic and associated changes in the landscape, which led to the extinction of the North American and numerous other rhino representatives in the original range. Towards the end of the Pleistocene there was another phase of extinction, during which all northern Eurasian representatives disappeared. The white rhinoceros and black rhinoceros that still exist today have survived in sub-Saharan Africa, as well as the armored , Java and Sumatran rhinoceros in south to south-east Asia , some of which have once again shrunk considerably due to the destruction of their habitat and poaching .

features

The Sumatran rhinoceros , the smallest and most primitive recent rhinoceros species
The endangered Java rhinoceros at London Zoo (1880)
Black rhinoceros in the Ngorongoro Crater

Rhinos are large to very large mammals. In the representatives living today, they have a head-torso length between 2.5 and 3.8 m (plus a 40 to 60 cm long tail) with shoulder heights varying from 1.2 to 1.8 m and a body weight fluctuating between 500 and 3,600 kg. The largest rhinoceros species living today is the white rhinoceros ( Ceratotherium simum ). However, fossil species were found to be significantly larger. Some representatives of Elasmotherium and Brachypotherium each reached a body weight of more than 5,000 kg.

In general, rhinos have a massive body with a large head and short, strong legs. Each foot has three toes that each end in wide hooves. The skin is thick and gray or brown in color. In the Asian species , the skin at the base of the neck and legs is heavily folded, so that it looks as if the animals are armored. Most rhinos today are hairless with the exception of the tips of their ears and tails. However, the Sumatran rhinoceros ( Dicerorhinus sumatrensis ) still has a fairly thick coat of hair . The fossil woolly rhinoceros ( Coelodonta antiquitatis ) has been shown to have a very dense fur from carcasses preserved in the permafrost of Siberia ; it is sometimes assumed for other extinct rhino species. In addition, rhinos have poor eyesight, but this disadvantage is offset by a keen sense of smell and very good hearing .

Skull and dentition features

Rhinos have a very large skull, which is usually quite elongated and erect on the occiput . The nasal bone is often massive and clearly arched forward and protrudes over the intermaxillary bone . At the points where the horns start, there are clear, mostly pearl or cauliflower-shaped roughened structures on the bone surfaces. The brain capsule is relatively small.

The set of teeth is very different, but is generally reduced in today's rhinos. The entire front dentition consisting of incisors and canines is missing in the African rhinos . The Asian rhinoceros, on the other hand, still have one or two pairs of incisors per lower and upper half of the dentition. The innermost upper incisors (I1) have a chisel-like shape, the second lower incisors (i2) are reminiscent of tusks or small tusks. This is known as the "chisel-tusk" complex. It is also found in many extinct representatives and used to distinguish the rhinos from their phylogenetically closest relatives. The molars have two as in all perissodactyla from enamel formed cleats on the occlusal surface ( bilophodont on). As a general characteristic, the enamel runs characteristic "π" -shaped on the chewing surfaces of the two anterior maxillary molars, while it is designed "L" -shaped on the lower jaw molars. This feature also applies to all fossil rhinos and their immediate relatives, so that this had already emerged about 50 million years ago. Depending on the diet, the back molars can have a low or high crown ( brachyodont or hypsodont ). The premolars are largely molarized in recent species, that is, they differ only slightly from the molars. However, depending on the phylogenetic development, there are different degrees of molarization of the premolars in the individual fossil species. Ancient rhinos still had the complete set of teeth consisting of three incisors, one canine, four premolars and three molars per jaw branch; the most reduced dentition was found in Elasmotherium with only two premolars and three molars per jaw branch. In addition, the molars of this genus of rhinoceros had the highest tooth crowns within the entire order of the odd ungulate , and they are surpassed in this characteristic only by a few representatives of rodents . The reduction in the number of teeth began very early in the tribal history of the rhinos. As a rule, the lower canine receded first, followed by the outermost lower and upper incisors and the upper canine.

horns

Forest rhinoceros skull ; the cauliflower-like roughened areas on the bone surface mark the attachment points of the nasal and frontal horns
Indian rhinoceros in Kaziranga National Park

An essential, visually distinctive and eponymous feature of the rhinos are the horns. The recent representatives have one horn or two horns depending on the species. The Asian armored rhinoceros ( Rhinoceros unicornis ) and the Java rhinoceros ( Rhinoceros sondaicus ) have only one anterior horn (nasal horn) which grows out of the nasal bone. The African rhinoceros white and black rhinoceros ( Diceros bicornis ) and the Asian Sumatran rhinoceros, on the other hand, have two horns, with the nasal horn also sitting on the nose, but the rear (frontal horn) growing on the frontal bone . In the case of extinct rhinoceros, the horns can usually only be detected by means of the attachment points on the skull. For example, members of the genus Elasmotherium had only one probably very large horn on their forehead, while those from Diceratherium had two nasal horns and those from Aceratherium probably had none at all.

The horn consists of agglutinated keratin , a fibrillar protein that is also found in hair , and despite its strength, contains neither bone substance nor, as is sometimes erroneously claimed, ivory . It is composed of numerous long, thread-like strands, called horn columns or filaments, the spaces between which are consolidated with horny substance . These threads run through the entire length of the horn, but taper significantly towards the top. The core of the horn is much more solid and mostly colored black, towards the outside it becomes much more fibrous and takes on a light gray color. Fossil horn has so far only been handed down from the woolly rhinoceros, but basically has the same structure as the recent one. The horn wears out continuously over time by rubbing against the ground or stones , it can also break off during a fight with fellow dogs or as a result of traumatic experiences, but it grows again throughout life. The largest known horn to date measured 1.58 m over the front curve.

Internal organs

Like all odd ungulates, rhinos are good rectal fermenters , so digestion takes place largely in the intestines . The stomach reaches a length of 120 cm. The appendix is between 60 and 90 cm long, the colon 5 to 8 m, the entire intestinal tract can be up to 20 m long. Microorganisms in the rearmost part of the intestine are used to break down indigestible parts of plants . The heart weighs up to 5 kg. The males do not have a scrotum ; the testicles are inside the body.

distribution and habitat

Distribution of today's rhinos in South Africa , Central Africa and Asia , present and historical; originally the rhinos were spread over Eurasia, Africa and North America

Today rhinos live in sub-Saharan Africa and in South and Southeast Asia both in savannah landscapes and in tropical rainforests in high and lowlands. The original distribution was much wider. The genetically oldest species can be found in Eurasia and North America in the middle Eocene around 50 million years ago . The rhinos first reached Africa with the closure of the Tethys Ocean and the creation of a land bridge in the early Miocene around 20 million years ago. At the end of the Miocene and the transition to the Pliocene 4 million years ago, the rhinos became extinct due to climate changes in North America.

During the Pleistocene, with its strongly fluctuating climate, rhinos were also widespread in the far north of Eurasia and lived in subarctic tundra landscapes . At the end of the last ice age , the rhinos disappeared from North Asia and Europe. In the course of their tribal history , the different rhinoceros species had occupied almost all ecological niches accessible to large, terrestrial mammals.

Way of life

Social behavior, reproduction and development

Indian rhinoceroses : mother with young in Chitwan National Park

Rhinos often live as solitary animals, but they can also appear in small, matriarchal herds in savannahs . Bulls are mostly loners and live territorial. The individual animals inhabit narrowly defined territories , which are marked with urine and feces , as well as the paths that are frequently used. During the day rhinos sleep or hang out in the wallowing areas, and you can see them actively eating at dusk and at night. They are shy animals that avoid human proximity. Reports of the animals' aggressiveness are usually greatly exaggerated. When it comes to attacks, the attacks are hardly targeted, but can result in fatal injuries due to the horns, the front teeth and the strength and mass of the animal. A rhinoceros also runs at speeds of up to 45 km / h (12.5 m / s), which is just about better than top human athletes. It can change direction abruptly. In addition, two different strategies can be observed in fighting. The Asian rhinoceros, all of which have front teeth, only rarely use their mostly small horns in fights, which are then fought in a highly ritualized manner in horn fights. The actual weapons are the dagger-like incisors of the lower jaw, with which they tear dangerous and deep wounds. The African rhinoceros, which do not have a front dentition, use their often much longer horns - especially the nasal horn - in addition to threatening gestures, also actively as a weapon to defend themselves, territory and also for food, in order to weaken the opponent by impaling.

During the rutting season of a cow there can be fights among the bulls, with the winner vying for the female in a striking way. This takes place through mutual hunting or mock fights, then copulation occurs . After a gestation period of 15 to 18 months, a young is born that can stay with the mother for two and a half to three years. If a second young animal is born, the older one is chased away by the mother, at least for the period of suckling. The life expectancy of today's rhinos is between 30 and a maximum of 50 years. The age of fossil species is mainly determined in comparison to the white rhinoceros and is based on anatomical features such as tooth eruption , degree of chewing of the individual teeth or stages of adhesion of certain bone sutures . In general, life expectancy in mammals is closely related to the body weight of the adult animals; in extinct rhinos, which were similar in size to today's, it was probably in an equivalent range. More detailed analyzes of individual fossil populations are only rarely available, for Pleistocene representatives this largely affects the woolly rhinoceros ( Coelodonta antiquitatis ), in older forms those for the more common genera such as Teleoceras or Chilotherium from the Miocene are known.

The rhinoceros species living today, which inhabit largely open landscapes, such as the African rhinoceros and the Indian rhinoceros, are often accompanied by birds such as maggot cutters or cattle egrets , which sit on their skin and cleanse them of parasites. The alarm calls of the maggot chopper also warn the animals of dangers or disturbances, including humans, at distances of 27 to 61 m. With a higher number of maggot cutters, an animal can locate the potential disturbance at a greater distance, but the calls do not provide any information about the direction. The rhinos determine this through their sense of smell. This has not yet been observed in the Java and Sumatran rhinos, which predominantly live in the tropical rainforest, which is related to their way of life in closed landscape areas. Young animals can in individual cases be captured by big cats and hyenas , occasionally also by wild dogs , adult rhinos have no natural enemies apart from humans. Most of the fossil representatives also rarely had to fear confrontations with predators, but occasionally bite wounds occur in representatives that are now extinct, some of which were caused by very large predators .

nutrition

Grazing white rhinoceros

All rhinos feed exclusively on vegetable food and are adapted to this diet with broad molars. The species have specialized in different plant foods. Four of the five rhinoceros living today prefer soft, plant-based diets, such as leaves , branches , twigs, buds and fruits ( browsing ). The molars of these types usually have low crowns and less dental cement. In addition, these rhinoceros have a rather rectangular short occiput due to the high head posture. The Indian rhinoceros ( Rhinoceros unicornis ) also feeds on grasses , but like the other rhinoceros that feed on leaves, it has a pointed, flexible upper lip. The white rhinoceros is the only recent rhinoceros species that has been completely adapted to grass forage ( grazing ). Since grasses contain silicic acid , which is very hard, teeth with high crowns and a high proportion of dental cement have formed due to the high abrasion when chewing. The long-term low head posture while eating led to an evolutionary lengthening of the occiput and thus to a low head posture and the formation of a neck hump. Furthermore, the white rhinoceros has eponymous broad, bulging lips.

Much of the fossil rhinos specialized in soft plant food because of their anatomy. Nevertheless, during the history of the tribe, as a result of climate and associated changes in the landscape within the various rhinoceros lines, independently of one another ( convergent evolution ), specialization in grass food came about. In the more modern rhinoceroses of the Rhinocerotinae group , this was associated with changes in the anatomy of the skull, including the elongation of the occiput and a resulting, low-hanging head. This mainly happened towards the end of the Miocene and during the Plio and Pleistocene. In addition to the white rhinoceros living today, the most famous fossil grass-eating representatives are the woolly rhinoceros, the steppe rhinoceros ( Stephanorhinus hemitoechus ) and the various members of the genus Elasmotherium . At least in the woolly rhinoceros, the fossilized ice mummies have proven that the mouth is shaped like the white rhinoceros. Another adaptation model of grass food is in phylogenetically older lines as the Aceratheriinae detectable. Here the limbs were shortened, causing the body and head to be low. These rhinoceros representatives could reach the ground with only a slight lowering of the head; the occiput was not lengthened, so that the skull was carried mainly horizontally. Well-known genera here are Chilotherium and Teleoceras from the late Miocene. However, both development models are characterized by the increase in the tooth crowns in order to counteract increased abrasion from the hard grass forage.

Systematics

External system

Live reconstruction of the early rhinoceros relative Paraceratherium
Rhino related cladogram according to Wang et al. 2016
 Rhinocerotoidea  

 Hyracodontidae (†) 


   


 Amynodontidae (†) 


   

 Indricotheriidae (= Paraceratheriidae †) 



   

 Eggysodontidae (†) 


   

 Rhinocerotidae 





Template: Klade / Maintenance / Style

The closest relatives of the rhinos living today are the horses and tapirs . The separation from the tapirs took place about 47 to 57 million years ago, the horses had split off 56 to 60 million years ago. The rhinos are part of the superfamily Rhinocerotoidea (rhinoceros). In a classic view, two other extinct families belong to this: the Amynodontidae and the Hyracodontidae , the latter forming a more closely related group together with the rhinos, which differ from the Amynodontidae by a longer molar row compared to the skull and in individual tooth features. The main differences can be found in the front dentition structure. The rhinos have a typical "chisel-tusk" formation of the upper and lower incisors, the Amynodontidae have enlarged canines and small incisors, while the Hyracodontidae show a more diverse pattern. The Amynodontidae existed from the Middle Eocene to the Middle Miocene and occurred in Eurasia and North America. Representatives of these rhinoceros were sometimes as big as today's rhinos, but did not wear a horn. They lived semi- aquatic life and probably ate aquatic plants . The Hyracodontidae, which also occurred from the Middle Eocene to the Early Miocene, show a high degree of variability in this definition, which is expressed in three existing subfamilies. These include the relatively small Hyracodontinae, the Allaceropinae and the large Indricotheriinae. With Paraceratherium (also known by the synonyms Baluchitherium and Indricotherium ), the latter represented the largest known land mammal in geological history. It was long-necked and hornless and mainly occurred in the Oligocene in Asia . Newer studies consider the Hyracodontidae as a non-uniform group. They are therefore split into the Hyracodontidae in the narrower sense, the Eggysodontidae (roughly equivalent to the Allaceropinae) and the Indricotheriidae (= Paraceratheriidae). The latter two are much closer to the rhinos with their more specialized anterior teeth than the classic Hyracodontidae with their more generalized incisors and canines.

Internal system

Internal classification of rhinos according to Heissig et al. 2007 and Prothero et al. 1986, 1989
 Rhinocerotidae  
  Diceratheriinae  

 Diceratheriini (†)


   

 Trigoniadini (†)



   
  Aceratheriinae  

 Aceratheriini (†)


   

 Teleoceratini (†)



  Rhinocerotinae  

 Menoceratini (†) 


   

 Elasmotheriini (†)


   

 Rhinocerotini 






Template: Klade / Maintenance / Style

The actual rhinoceros (Rhinocerotidae) developed several lines: The subfamily of the Diceratheriinae includes two tribes with the Diceratheriini and the Trigoniadini . They were characterized by paired horns on their noses and were the first rhinos with such formations. They lived largely in the Oligocene and Miocene, mainly in North America , where they were the largest endemic group of mammals until 21 million years ago. In Eurasia , fossils of this rhinoceros group are rarely known. Another subfamily are the Aceratheriinae , which only rarely had horns and are divided into the aceratheriini and teleoceratini tribes . Their development began as early as the Oligocene, but they partially survived until the end of the Pliocene and colonized Eurasia and North America as well as Africa .

The Rhinocerotinae subfamily also split into several lines. The tribe of the Menoceratini includes the trunk group, but was originally placed in the vicinity of the Diceratheriinae due to the paired nasal horns. But they have much more modern development features than these. The Elasmotheriini developed in the early Miocene, whereby the best known genus, the elephant-sized Elasmotherium , still lived in the last glacial period ( Weichselian glacial period ) and was characterized by an oversized horn, possibly up to 2 m long.

Relationship of rhinos based on genetic data from Margaryan et al. 2020
  Rhinocerotinae  
  Elasmotheriini 

 Elasmotherium (†)


  Rhinocerotini 
  Dicerorhinina 


 Stephanorhinus (†)


   

 Coelodonta (†)



   

 Dicerorhinus



   
  Dicerotina 

 Ceratotherium


   

 Diceros



  Rhinocerotina 

 Rhinoceros





Template: Klade / Maintenance / Style

The five species living today all belong to the rhinocerotini tribe. Your closest relatives are probably the Elasmotheriini. According to molecular genetic studies, there is a deep temporal gap between the two lines , since their separation began in the Paleogene ; depending on the study, the data range from the Middle Eocene around 47 million years ago to the Lower Oligocene around 31 million years ago. The rhinocerotini can be divided into three groups. The division into today's two Asian and one African lines took place around 29 to 30 million years ago in the Lower Oligocene. The highly endangered Sumatran rhinoceros ( Dicerorhinus sumatrensis ) is the only surviving species of the primal group, the Dicerorhinina. The rhinoceros genera Coelodonta with the well-known woolly rhinoceros and Stephanorhinus , to which the less well-known forest rhinoceros belongs, are also assigned to the Dicerorhinina, which were distributed over large parts of northern Eurasia during the Pleistocene , but the genus Coelodonta split from the line of the 21 million years ago Sumatran rhinoceros. Another group separated from the same lineage 26 million years ago, the Rhinocerotina, which comprises two species: the endangered Indian rhinoceros ( Rhinoceros unicornis ) and the highly endangered Java rhinoceros ( Rhinoceros sondaicus ). However, their lines were only formed about 11.7 million years ago. The third group, the Dicerotina, includes the two African species, the white rhinoceros ( Ceratotherium simum ) and the black rhinoceros ( Diceros bicornis ), whose splitting began around 17 million years ago.

Alternative internal classification of rhinos according to Cerdeño 1995
 Rhinocerotidae  

 incertae sedis (†) 


   
  Rhinocerotinae  

 incertae sedis (†) 


  Rhinocerotini  

 Rhinocerotina 


   

 Dicerotina 


   

 Iranotheriina (†) 


   

 Elasmotheriina (†) 


Template: Klade / Maintenance / 3Template: Klade / Maintenance / 4


  Aceratheriinae  

 Aceratheriini (†) 


   

 Teleoceratini (†) 





Template: Klade / Maintenance / Style
Alternative internal classification of rhinos according to Antoine 2003
 Rhinocerotidae  

 incertae sedis (†) 


   
  Rhinocerotinae  
  Rhinocerotini  

 Rhinocerotina 


   

 Teleoceratina (†) 



   

 Aceratheriini (†) 



  Elasmotheriinae  

 Diceratheriini (†) 


  Elasmotheriini  

 Menoceratina (†) 


   

 Elasmotheriina (†) 






Template: Klade / Maintenance / Style

The rhinoceros system shown here is based on the work of Kurt Heissig , Donald R. Prothero and Colin Peter Groves . The systematic position of the living rhinoceros species as a tribe or sub-tribus is under discussion, but the position as sub-tribus seems to be justified. The position of Dicerorhinus is also problematic . which is sometimes assigned to the Rhinocerotina sub-tribus or no special sub-tribus.

There are also other classification schemes of rhinos such as that of Pierre-Olivier Antoine, in which the two subfamilies Elasmotheriinae and Rhinocerotinae are distinguished. The former includes the tribes Elasmotheriini and Diceratheriini, while Rhinocerotini and Aceratheriini belong to the Rhinocerothinae. An additional classification scheme also comes from Esperanza Cerdeño, which highlights the subfamilies Acertheriinae with the tribe Alicornopini or Teleoceratini and the subfamily Rhinocerothinae with the sole tribe Rhinocerotini.

Overview of the recent and fossil rhinoceros taxa

Including the recent representatives, around 60 genera with several hundred species have been described to date . The classification scheme used here largely follows Prothero and Schoch 1989, whereby changes to the structure from Heissig 2007 and further revisions and additions from more recent times have been taken into account:

Size and habitus comparison of recent and extinct rhino forms
  • Family: Rhinocerotidae Owen , 1845.

Tribal history

Adaptive radiation

The rhinoceros family is one of the most successful and diverse groups of mammals in recent geological history and can be traced back around 50 million years. During this time a broad evolutionary development and adaptation to the respective habitats took place ( adaptive radiation ). Representatives of the rhinos occupied almost every terrestrial biotope. Within the evolutionary history there have been numerous anatomical changes. General evolutionary trends in rhinos, which largely went through all lines and which each represent adaptations to certain biotopes , are the shortening and widening of the skull, especially in the anterior facial area, length reduction of the limbs, reduction of the teeth, molarization of the premolars and enlargement of the height of the tooth crowns at the molars. Furthermore, two general development models can be identified: hornless rhinos with largely intact or extensive anterior dentition with enlarged teeth and horn-bearing rhinos with greatly reduced or nonexistent anterior dentition.

Eocene

Trigonias from the late Eocene and Oligocene

The earliest known precursor of the rhinos was Hyrachyus from the mid-Eocene of North America and Europe. It was about the size of a modern German Shepherd and had complete mammalian teeth with hardly any molarized premolars. Due to the very basal position, this genus is placed depending on the view of the tapiroids or rhinocerotoids. The first true representative rhinoceros was also found in Teletaceras in the middle or late Eocene, but like all early rhinos had no horn formation. It lived in both North America and Asia and was also relatively small. In the late Eocene, the first larger rhinos appeared, such as the cattle-sized Trigonias .

Oligocene

The first rhinos in Europe appear with the large and hornless Ronzotherium for the first time in the early Oligocene, while representatives of Guixia spread in Asia . In the late Oligocene, Europe and Asia shared numerous common rhino lines. With Protaceratherium, a member of the Aceratheriini developed, which were characterized by a long, trunk-like snout and a receding nasal region and specialized in soft vegetable foods. Likewise, with Diaceratherium (not to be confused with Diceratherium ) and Brachydiceratherium, the first representatives of the hippopotamus-like, short - legged Teleoceratini, some of which lived in open forest landscapes and later also lived in savannahs, emerged, some of which were specialized grazers and developed very high-crowned molars.

In North America their own rhino lines formed, but overall the diversity of the rhinos there was much lower compared to Eurasia. Subhyracodon , which also appeared in the late Eocene, developed into the Diceratherium line. These represented the first rhinos with horn formations, but in this group they sat in pairs on the nose. With the appearance of Subhyracodon and Diceratherium in North America at the end of the Oligocene, all primitive rhinos with four-toed forefeet were replaced by more modern rhinos with three-toed forefeet, a process that lasted well into the Miocene in Eurasia and Africa .

Miocene

Teleoceras from the Miocene

In the Miocene there was a strong radiation of the rhinos with a great variety of forms. The rhinos first set foot on African soil in the early Miocene. Early finds are proven with Aceratherium and Brachypotherium in Egypt and Libya . Representatives of the Aceratheriini also reached North America and formed their own genera with Floridaceras and Aphelops , among others . The Teleoceratini also emigrated to North America. Here in Nebraska on Verdigre Creek in the Ash Hollow Formation , where a complete herd with intact stomach contents was buried under volcanic ash , one of the best evidence of the genus Teleoceras , which gave this rhinoceros its name and which was also one of the most dominant herbivores of that time in North America was. Furthermore, the Menoceras from Eurasia appeared , which like Diceratherium had paired nasal horns but is a basal representative of the Rhinocerotinae.

Various originally North American rhinos also immigrated to Eurasia, as did individual representatives of the Diceratheriini. What is important here, however, is the spread of the hornless rhinos, the Aceratheriini, which produced numerous forms and which, in addition to the aforementioned Aceratherium, also include Alicornops and Chilotherium . The latter in particular was very common. Furthermore, members of more modern rhino forms, such as Bugtirhinus as the basal form of the Elasmotheriini, can be detected for the first time in the early Miocene . The first appearance of the genera still existing today, such as the African forms Ceratotherium and Diceros in the late Miocene , is also significant . The distribution area of Diceros. originally it was much larger because the genealogically old species Diceros gansuensis. also occurred in East Asia 5 to 7 million ago. Furthermore, Dicerorhinus, early representatives of the Dicerorhinina have been found in Eurasia , while Sinotherium and Ningxiatherium represent clearly more developed members of the Elasmotheriini. The Rhinocerotina can be detected in this time phase with Gaindatherium and Punjabitherium .

At the end of the Miocene, numerous rhinoceros species became extinct due to climatic cooling combined with the spread of open steppe landscapes. This particularly affected the Aceratheriini and part of the Teleoceratini. In North America the entire rhinoceros fauna disappeared, in Eurasia only Elasmotheriini, Dicerorhinina and Rinocerotina survived. In Africa the Dicerotina and some species of Brachypotherium from the group of Teleoceratini also kept.

Plio and Pleistocene

Woolly rhinoceros from the late Pleistocene

The period of the Plio and Pleistocene was mainly characterized by the further development of modern rhinoceros groups such as Elasmotheriini and Rhinocerotini. The latter include the species that still exist today. Above all the primeval Dicerorhinina of Eurasia showed a great diversity. However, little is known of the direct ancestors of the Sumatran rhinoceros. The rise of the Coelodonta, known since the middle Pliocene , to the woolly rhinoceros that lived at the end of the Pleistocene is significant . The sister group Stephanorhinus split into several species adapted to different biotopes, as shown by the Middle Pleistocene forms of the forest and steppe rhinoceros. However, both rhino lines end in the late Pleistocene. The genus Rhinoceros has been detectable since the Pliocene and split into several forms. In northern Eurasia, especially in Central Asia , the various representatives of Elasmotherium have been handed down. This rhinoceros genus also died out in the Young Pleistocene. In Africa, Diceros and Ceratotherium develop into the species known today.

At the end of the Pleistocene there was another extinction wave, in the course of which the rhino lines of northern Eurasia disappeared (see Quaternary extinction waves ). Only the rhinoceros representatives who still exist today survived.

Taxonomy

Richard Owen , 1856

The name Rhinoceros was introduced by Linnaeus in 1758 as a scientific name for the rhinoceros, but the name has been used since at least the beginning of the 16th century, for example by Albrecht Dürer in his famous woodcut Rhinocerus , which he made in 1515. One of the earliest references to the use of the name comes with rhinókerôn from the Bibliothéke historiké of the ancient historian Diodor von Agyrion from the 1st century BC. The name is made up of the Greek words ῥίς ( rhīs "nose"; genitive rhinos ) and κέρας ( kéras "horn"), so it clearly refers to the formation of horns on the nose. Linnaeus distinguished the then known two types: R. unicornis , the rhinoceros , which he situate in India and Africa, and R. bicornis , the black rhino , which according to his opinion, but lived in India. Linnaeus' errors regarding distribution are probably based on the use of often older sources in the preparation of his work Systema Naturae . Numerous scholars of his time refused to divide rhinos into two types and instead preferred only one that showed certain variations. The contradiction only resolved when the Dutch anatomist Petrus Camper studied a two-horned rhinoceros from South Africa in 1771 and noticed not only the second horn but also differences in the structure of the teeth; however, his results were not fully published until 1782. Around the same period, 1769, the German naturalist Peter Simon Pallas discovered a two-horned rhinoceros in Siberia, the woolly rhinoceros .

The British naturalist John Edward Gray introduced the family name Rhynocerotidae in 1821. His brief description was: “Nose short, rounded, bones very thick, bearing a horn formed of agglutinated hair; toes three to each foot; stomach simple; intestine and caecum large "(" nose short, rounded, bones very thick, horn-bearing, consisting of coalesced hair; three toes on each foot; stomach simple; intestine and appendix large "). Due to the incorrect spelling, this name is not officially recognized, but Gray formally differentiated in his description the two genera Rhinoceros (Indian rhinoceros ) and Diceros (black rhinoceros) that still exist today . As early as 1811, the German zoologist Johann Karl Wilhelm Illiger had suggested the name Nasicornia for the rhinos, which, since it was not based on a generic name, is also invalid. In 1845 Richard Owen , next to Charles Darwin one of the most important natural historians of the Victorian era , presented a study in which he included the rhinoceros family and also considered the teeth that are important for mammalian taxonomy. Here he correctly referred to the rhinos as Rhinocerotidae, which is why many experts prefer the group name "Rhinocerotidae Owen , 1845" to the actually incorrect name "Rhinocerotidae Gray , 1821".

Rhinos and humans

Rhinos in art and culture

Depictions of animals in the Chauvet grotto (replica): horses (above), aurochs (left) and woolly rhinos (one in the middle, two below right)

In terms of body size and habitus , rhinos are among the most impressive land-living mammals and have therefore also found their way into human art and culture, this is particularly true of hunter-gatherer populations. The earliest known representations of rhinos can be found in the Western Eurasian Upper Paleolithic (40,000 to 10,000 years ago) and are at least 31,000 years old. The paintings of the Franco-Cantabrian cave art , where rhinos were depicted in more than half a dozen caves with over 80 representations, should be emphasized here . Most often they are preserved in the Grotto Chauvet ( France ) with 65 drawings, some in red or black color pigments, one panel alone contains 17 depictions of an animal in motion. Furthermore, in Europe outside of this cultural area in the mobile Upper Palaeolithic small art there are illustrations and representations of rhinos in the form of bone or stone carvings , but also clay figures modeled as small statuettes, which are among the oldest ceramic objects in the world. All of these artifacts are interpreted today as depictions of the woolly rhinoceros, which lived in the northern Eurasian cold steppes , some older ones sometimes also as the steppe rhinoceros. Some researchers believe that individual characteristic rhinoceros depictions, such as those in the Rouffignac Cave , could be viewed as images of Elasmotherium , but this genus was neither spatially nor temporally so widespread.

Also prehistoric hunter-gatherer communities in other parts of the world depicted rhinos. In South Asia there are numerous paintings in caves and abrises, which in this case depict the Indian rhinoceros. These are assigned to the Mesolithic there and are between 12,000 and 7,000 years old. Significant and among the oldest in this region are those from the Marodeo rock near Pachmarhi in the Indian state of Madhya Pradesh or a hunting scene, shown near Mirzapur in the Indian state of Uttar Pradesh . Numerous rock carvings and carvings of both white and black rhinoceros are known from large areas of southern Africa , which are predominantly assigned to the Khoisan population groups and can be up to 14,000 years old, but some are also significantly younger. These are sometimes very numerous, with more engravings than drawings being observed in some areas. In the South African provinces of North Cape, Northwest and Free State alone , more than 500 depictions have survived.

The "Golden Rhinoceros" from Mapungubwe ; 1075-1220; Mapungubwe Collection (University of Pretoria Museums).

With the settling down of humans, the images of rhinos are rapidly declining. Representations of animals on seals of the Indus culture are known from the Copper Age , and there are also isolated reliefs in ancient Egypt . Relief figures of one-horned rhinos on temple friezes in Angkor Vat ( Cambodia ), which date from the 12th century and are among the few images of the Java rhinoceros due to their historical range, are also remarkable. The "Golden Rhinoceros" from the "Original Gold Grave" on Mapungubwe Hill in the South African province of Limpopo is of great cultural importance for southern Africa . The archaeological site complex was created by local tribes in the 11th to 13th centuries and excavated in the 1920s to 1930s. The small rhinoceros figurine with a height of 5.5 cm possibly formed, together with other zoomorphic statuettes such as a buffalo, a big cat, an elephant and a crocodile, a decorative element of a larger vessel that was probably used in fortune telling . For European art history outstanding of widespread woodblock are Rhinocerus by Albrecht Durer from 1515, produced in only indirect reports of an Indian rhinoceros in Portugal, and the oil painting Rhinoceros Clara by Jean-Baptiste Oudry from the year 1749th

Threat and protection

Population estimates of the five recent rhino species
Art Stock size 2007 Stock size 2012 Stock size 2018
White rhinoceros 14,500 20,400 18,067
Black rhinoceros 3,725 5,050 5,495
Indian rhinoceros 2,619 3,300 3,588
Sumatran rhinoceros 275 <100 40-78
Java rhinoceros 55 35-44 65-68
The white rhinoceros bull Tsavo in the circus

The rhinos living today are non- domesticable wild animals due to several factors - enormous size, slow growth, territorial loners, few offspring in human care . As a result, the importance of the animal group for today's humans as a supplier of food and raw materials is relatively minor, but there is a great demand for rhinoceros horns in Asia . These are traditionally used in the Middle East , especially in northern Yemen , for the handles of the Jambia dagger, which serves as a status symbol. In East Asia, however, they are part of handicraft carvings and traditional Chinese medicine . In the latter, the horns are used, especially in powdered form, as a medicine against fever and pain, an often assumed use as an aphrodisiac is not historically guaranteed. Mainly because of this market, rhinos are threatened with extinction from the associated poaching . In 2017, up to 45,000 US dollars for every kilogram of horn of an Asian rhinoceros was traded on the black market, and up to 10,000 US dollars in 2011 for the corresponding amount of an African rhinoceros. Due to the high demand for horn from East Asia, not only has the number of wild rhinos killed by poaching, especially in southern Africa, increased, but there has also been an increase in thefts in museums , collections and auction houses and the associated smuggling . However, several tests carried out by the pharmaceutical industry and life science research institutes did not reveal any medical effects. However, there were always rumors of miraculous cures, for example, the demand last increased in 2009 after a Vietnamese man claimed to have conquered his cancer by taking horn. Other causes of the threat to today's rhino species are the destruction of habitats by agriculture or the construction of traffic routes, but also the expansion of human settlements to the limits of the protected areas.

Recently, however, there has been a slight recovery in the populations of some species. At the end of 2010, around 20,000 white rhinos and over 5,000 black rhinos were living in Africa again. Both stocks have almost doubled since 1995. A large proportion of the rhinos live on it in South Africa, mostly in fenced and guarded reserves. In Kenya there are also various private and public protected areas, e.g. As the private farm Ol ari Nyiro of Kuki Gallman or the Lake Nakuru National Park . According to the IUCN , the total population of African rhinos has stabilized, but individual subspecies are in some cases severely threatened. The northern white rhinoceros ( C. s. Cottoni ), which only contains a few specimens and is to be saved from extinction through a breeding program in the Ol Pejeta reserve in Kenya, is particularly critical. Due to the increasing number of rhinos killed by poachers in southern Africa (almost 800 rhinos in 2013), new countermeasures were introduced in 2013 in addition to armed rangers, dehorning of wild animals or the relocation of individual individuals or smaller populations. These include, among other things, the injection of antiparasitic drugs into the horns, mainly drugs against external parasites , which are harmless to rhinos in normal use, but cause nausea or convulsion in humans and thus make the horns unusable for the traditional Chinese medicine market. Another method is to mark the horns with dyes that color their insides red or pink. This should help to track the international trade in illegally captured horns, as these can be detected by scanners at airports , comparable to banknotes marked by a similar process . So far, environmentalists have been rather skeptical about these approaches, on the one hand because a few rhinos died while carrying out the procedure, and on the other hand because hardly any scientific studies were carried out on the new measures and the associated potential health risks for the treated animals. Furthermore, they can only pretend that the rhinoceros populations are more secure, since poaching has declined in individual protected areas with animals treated in this way, but the majority of the animals had abandoned them due to the pressure of poaching. A shift in hunting pressure cannot be ruled out. In March 2014, WWF Germany reported about a project in which a chip is to be implanted in 1000 rhinos to be protected in Kenya in order to better catch poachers.

Female Sumatran rhinoceros (“Rosa”) in the Sumatran Rhino Sanctuary in Sumatra

Protection efforts by India and Nepal were also successful, so that the population of Indian rhinos rose to 2,850 animals at the end of 2010, which is a significant increase compared to 1995; at present it is assumed that there are more than 3,500 animals. The population of Sumatran rhinos, on the other hand, has declined from a little more than 300 to around 220 to 280 animals in the same period, and in some cases fewer than 100 animals are accepted - the reason is obviously that the governments of Indonesia and Malaysia have only limited financial resources for the Provide protection for these animals. The Java rhinoceros is now the most endangered large mammal on earth. An estimated 60 specimens inhabit remnants of the former range of the species in the west of Java . The last remaining small population of the Java rhinoceros in Vietnam was declared extinct by the WWF in October 2011 . Thus the Vietnamese subspecies R. s applies . annamiticus of the Java rhinoceros officially as extinct. Due to the high risk of isolating the last cows capable of reproduction and the fact that rhinos generally rarely reproduce in human care, breeding programs are difficult to implement, so that earlier conservation efforts have resulted in the creation of new protected areas and the rescue of the Restrict residual populations. Two major recent projects in this context, the started to protect the Sumatran rhino breeding program in 1997 in the Sumatran Rhino Sanctuary in Sumatra and the 2011 project launched Javan Rhino Study and Conservation Area for the conservation of the Javan Rhino.

literature

  • Anonymous (ed.): The rhinos: encounter with primeval colossi. Fürth, Filander Verlag, 1997, ISBN 3-930831-06-6 .
  • Esperanza Cerdeño: Diversity and evolutionary trends of the family Rhinocerotidae (Perissodactyla). In: Palaeo. 141, 1998, pp. 13-34.
  • Claude Guérin: La famille des Rhinocerotidae (Mammalia, Perissodactyla): systématique, histoire, évolution, paleoécologie. In: Cranium. 6, 1989, pp. 3-14.
  • Kurt Heissig: The Rhinocerotidae. In: Donald R. Prothero, RM Schoch (Ed.): The evolution of the Perissodactyls. New York 1989, pp. 399-417.
  • Donald R. Prothero, Claude Guérin, Earl Manning: The history of Rhinocerotoidea . In: Donald R. Prothero, RM Schoch (Ed.): The evolution of the Perissodactyls. New York 1989, pp. 321-340.
  • Donald R. Prothero: Fifty million years of rhinoceros evolution. In: OA Ryder (Ed.): Rhinoceros biology and conservation: Proceedings of an international conference, San Diego, USA Zoological Society, San Diego 1993, pp. 82-91.
  • Rudolf Schenkel, Ernst M. Lang : The behavior of the rhinos (= Handbook of Zoology. 8 (46)). 1969, pp. 1-56, ISBN 3-11-000664-2 .

Individual evidence

  1. ↑ a b c d e f g h i Esperanza Cerdeño: Diversity and evolutionary trends of the family Rhinocerotidae (Perissodactyla). In: Palaeo. 141, 1998, pp. 13-34.
  2. ↑ a b Ralf-Dietrich Kahlke, Frédéric Lacombat: The earliest immigration of woolly rhinoceros (Coelodonta tologoijensis, Rhinocerotidae, Mammalia) into Europe and its adaptive evolution in Palaearctic cold stage mammal faunas. In: Quaternary Science Reviews. 27, 2008, pp. 1951-1961.
  3. ↑ a b c d e f g R. Fulconis: Save the rhinos: EAZA Rhino Campaign 2005/6. Info Pack, London 2005.
  4. ↑ a b c d e f g Kurt Heissig: The Rhinocerotidae. In: Donald R. Prothero, RM Schoch (Ed.): The evolution of the Perissodactyls. New York 1989, pp. 399-417.
  5. ^ Donald R. Prothero: Rhino giants. The paleobiology of Indricotheres. Indiana University Press, 2013, pp. 54-55, ISBN 978-0-253-00819-0 .
  6. ↑ Frederic Lacombat: The Evolution of the rhinoceros. In: R. Fulconis (Ed.): Save the rhinos: EAZA Rhino Campaign 2005/6. Info Pack, London 2005, pp. 46-49.
  7. ↑ a b Vladimir Zhegallo, Nikolay Kalandatze, Andrey Shapovalov, Zoya Bessudnova, Natalia Noskova, Ekaterina Tesakova: On the fossile rhinoceros Elasmotherium (Including the collections of the Russian Academy of sciences). In: Cranium. 22 (1), 2005, pp. 17-40.
  8. ↑ Jérémy Tissier, Pierre-Olivier Antoine and Damien Becker: New material of Epiaceratherium and a new species of Mesaceratherium clear up the phylogeny of early Rhinocerotidae (Perissodactyla). In: Royal Society Open Science. 7, 2020, p. 200633, doi: 10.1098 / rsos.200633 .
  9. ^ A b c d e f Donald R. Prothero, Claude Guérin, Earl Manning: The history of Rhinocerotoidea. In: Donald R. Prothero, RM Schoch (Ed.): The evolution of the Perissodactyls. New York 1989, pp. 321-340.
  10. ↑ Peter Kemnitz, Wolfgang Puschmann, Michael Schröpel, Dieter Krause, Rüdiger Schöning: Fine tissue studies on the structure and ontogenesis of the horn of rhinoceros, Rhinocerotidae: an atlas with new views on and about an old problem. In: Zoological Garden. 61 (3), 1991, pp. 177-199.
  11. ↑ ML Ryder: Structure of rhinoceros horn. In: Nature. 193, 1962, pp. 1199-1201.
  12. ↑ Fedor K. Shidlovskiy, Irina V. Kirillova, John Wood: Horns of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799) in the Ice Age Museum collection (Moscow, Russia). In: Quaternary International. 2011, pp. 1–5.
  13. ↑ О. Ф. Чернова, И. В. Кириллова: Новые данные о морфологии рога шерстистого носорога (Coelodonta antiquitatis BLUMENBACH, 1799). In: Труды Зоологического института РАН. 314 (3), 2010, pp. 333-342.
  14. ↑ Colin P. Groves: Species characters in rhinoceros horns. In: Journal of Mammals. 36 (4), 1971, pp. 238-252.
  15. ↑ Colin P. Groves, Fred Kurt: Dicerorhinus sumatrensis. In: Mammalian Species. 21, 1972, pp. 1-6.
  16. ^ WA Laurie, EM Lang, Colin P. Groves: Rhinoceros unicornis. In: Mammalian Species. 211, 1983, pp. 1-6.
  17. ↑ Nan E. Schaffer, George L. Foley, Sophia Gill, C. Earl Pope: Clinical implications of rhinoceros reproductive tractanatomy and histology. In: Journal of Zoo and Wildlife Medicine. 32 (1), 2001, pp. 31-46.
  18. ↑ a b c d e f Donald R. Prothero: Fifty million years of rhinoceros evolution. In: OA Ryder (Ed.): Rhinoceros biology and conservation: Proceedings of an international conference, San Diego, USA Zoological Society, San Diego 1993, pp. 82-91.
  19. ↑ a b Toni Bürgin, Urs Oberli: A chance find from the quarry. In: fossils. Magazine for collectors and amateur palaeontologists. 27 (4), 2010, p. 238 f.
  20. ↑ a b c Rudolf Schenkel, Ernst M. Lang: The behavior of the rhinos. (= Handbook for Zoology. 8 (46)). 1969, pp. 1-56, ISBN 3-11-000664-2 .
  21. ^ Adrian Lister and Paul Bahn: Mammuts - The Giants of the Ice Age. Sigmaringen, 1997 (p. 89).
  22. ↑ Nina V. Garutt: Dental ontogeny of the woolly rhinoceros Coelodonta antiquitatis (Blumenbach, 1799). In: Cranium. 11, 1994, pp. 37-48.
  23. ↑ Irina V. Kirillova, Fedor K. Shidlovskiy: Estimation of individual age and season of death in woolly rhinoceros, Coelodonta antiquitatis (Blumenbach, 1799), from Sakha-Yakutia, Russia. In: Quaternary Science Reviews. 29, 2010, pp. 3106-3114.
  24. ^ Matthew C. Mihlbachler: Demography of late Miocene rhinoceroses (Teleoceras proterum and Aphelops malacorhinus) from Florida: linking mortality and sociality in fossil assemblages. In: Paleobiology. 29 (3), 2003, pp. 412-428.
  25. ↑ Liang Zhong, Deng Tao: Age structure and habitat of the Rhinoceros Chilotherium during the Late Miocene in the Linxia Basin, Gansu, China. In: Vertebrata Palasiatica. 43 (3), 2005, pp. 319-230.
  26. ^ Roan D. Plotz, Wayne L. Linklater: Oxpeckers Help Rhinos Evade Humans. In: Current Biology. 30 (10), 2020, pp. 1965–1969.e2, doi: 10.1016 / j.cub.2020.03.015
  27. ↑ Nina Garutt: Traumatic skull damages in the woolly rhinoceros, Coelodonta antiquitatis BLUMENBACH, 1799. In: Cranium. 14, 1997, pp. 37-46.
  28. ↑ a b Deng Tao, Tseng Zhijie J: Osteological evidence for predatory behavior of the giant percrocutid (Dinocrocuta gigantea) as an active hunter. In: Chinese Science Bulletin. 55 (17), 2010, pp. 1790-1794.
  29. ↑ Jan van der Made, René Grube: The rhinoceroses from Neumark-Nord and their nutrition. In: Harald Meller (Hrsg.): Elefantenreich - Eine Fossilwelt in Europa. Halle / Saale 2010, pp. 382–394.
  30. ↑ Friedrich E. Zeuner: The relationships between skull shape and way of life in recent and fossil rhinos. In: Reports of the Natural Research Society in Freiburg. 34, 1934, pp. 21-80.
  31. ↑ Kurt Heissig: Family Rhinocerotidae. In: Gertrud E. Rössner, Kurt Heissig: The Miocene land mammals of Europe. Munich 1999, pp. 175-188.
  32. ↑ a b Haibing Wang, Bin Bai, Jin Meng, Yuanqing Wang: Earliest known unequivocal rhinocerotoid sheds new light on the origin of Giant Rhinos and phylogeny of early rhinocerotoids. In: Scientific Reports. 6 (39607), 2016, pp. 1-9, doi: 10.1038 / srep39607 .
  33. ↑ a b c d Christelle Tougard, Thomas Delefosse, Catherine Hänni and Claudine Montgelard: Phylogenetic Relationships of the Five Extant Rhinoceros Species (Rhinocerotidae, Perissodactyla) Based on Mitochondrial Cytochrome b and 12S rRNA Genes. In: Molecular Phylogenetics and Evolution. 19, 2001, pp. 34-44.
  34. ↑ Cynthia C. Steiner, Oliver A. Ryder: Molecular phylogeny and evolution of the Perissodactyla. In: Zoological Journal of the Linnean Society. 163, 2011, pp. 1289-1303.
  35. ↑ a b Pavel Kosintsev, Kieren J. Mitchell, Thibaut Devièse, Johannes van der Plicht, Margot Kuitems, Ekaterina Petrova, Alexei Tikhonov, Thomas Higham, Daniel Comeskey, Chris Turney, Alan Cooper, Thijs van Kolfschoten, Anthony J. Stuart, Adrian M. Lister: Evolution and extinction of the giant rhinoceros Elasmotherium sibiricum sheds light on late Quaternary megafaunal extinctions. In: Nature Ecology & Evolution. 3, 2019, pp. 31-38, doi: 10.1038 / s41559-018-0722-0 .
  36. ↑ a b c Donald R. Prothero: Rhinocerotidae. In: Christine M Janus, Kathleen M Scott, Louis L Jacobs (Eds.): Evolution of Tertiary mammals from North America. Vol. 1, Cambridge 1998, pp. 595-605.
  37. ^ Leonard B. Radinsky: The families of the Rhinocerotoidea (Mammalia, Perissodactyla). In: Journal of Mammalogy. 47 (4), 1966, pp. 631-639.
  38. ↑ a b c d Donald R. Prothero, Earl Manning, C. Bruce Hanson: The phylogeny of the rhinocerotoidea (Mammalia, Perissodactyla). In: Zoological Journal of the Linnean Society. 87, 1986, pp. 341-366.
  39. ↑ Bin Bai, Jin Meng, Yuan Qing Wang, Hai Bing Wang, Luke T. Holbrook: Osteology of the Middle Eocene ceratomorph Hyrachyus modestus (Mammalia, Perissodactyla). In: Bulletin of the American Museum of Natural History. 413, 2017, pp. 1-68.
  40. ^ A b c d e Kurt Heissig, Oldřich Fejfar: The fossil rhinos (Mammalia, Rhinocerotidae) from the Lower Miocene of Tuchorice in northwestern Bohemia. Sborník Národního Muzea v Praze. In: Acta Musei Nationalis Pragae (series B, Natural History). 63 (1), 2007, pp. 19-64.
  41. ^ A b c Donald R. Prothero, Robert M. Schoch: Classification of the Perissodactyla. In: Donald R. Prothero, RM Schoch (Ed.): The evolution of the Perissodactyls. New York 1989, pp. 530-537.
  42. ^ A b Pierre-Olivier Antoine, Francisco Alférez, Carlos Iñigo: A new elasmotheriine (Mammalia, Rhinocerotidae) from the early Miocene of Spain. In: Comptes Rendus Pale. vol 1, 2002, pp. 19-26.
  43. ↑ a b Ashot Margaryan, Mikkel-Holger S. Sinding, Shanlin Liu, Filipe Garrett Vieira, Yvonne L. Chan, Senthilvel KSS Nathan, Yoshan Moodley, Michael W. Bruford, M. Thomas P. Gilbert: Recent mitochondrial lineage extinction in the critically endangered Javan rhinoceros. In: Zoological Journal of the Linnean Society. , 2020, doi: 10.1093 / zoolinnean / zlaa004 / 5802322 .
  44. ^ Claude Guérin: La famille des Rhinocerotidae (Mammalia, Perissodactyla): systématique, histoire, évolution, paleoécologie. In: Cranium. 6, 1989, pp. 3-14.
  45. ^ Ludovic Orlando, Jennifer A. Leonard, Aurélie Thenot, Vincent Laudet, Claude Guerin, Catherine Hänni: Ancient DNA analysis reveals woolly rhino evolutionary relationships. In: Molecular Phylogenetics and Evolution. 28, 2003, pp. 485-499.
  46. ↑ Xiufeng Xu, Axel Janke, Ulfur Arnason: The Complete Mitochondrial DNA Sequence of the Greater Indian Rhinoceros, Rhinoceros unicornis, and the Phylogenetic Relationship Among Carnivora, Perissodactyla, and Artiodactyla (+ Cetacea). In: Molecular Biology and Evolution. 13 (9), 1996, pp. 1167-1173.
  47. ↑ a b Esperanza Cerdeño: Cladistic analysis of the family Rhinocerotidae (Perissodactyla). In: American Museum Novitates. 3143, 1995, pp. 1-25.
  48. ↑ a b c Pierre-Olivier Antoine: Middle Miocene elasmotheriine Rhinocerotidae from China and Mongolia: taxonomic revision and phylogenetic relationships. In: The Norwegian Academy of Science and Letters - Zoologica Scripta. 32, 2003, pp. 5-118.
  49. ↑ Colin P. Groves: The Rhinos - Tribal History and Kinship. In: The rhinos: encounter with primeval colossi. Fürth 1997, pp. 14-32, ISBN 3-930831-06-6 .
  50. ↑ Colin P. Groves: Rhinoceros sondaicus. In: Mammalian Species. 43 (887), 2011, pp. 190-208.
  51. ↑ Deng Tao: A primitive species of Chilotherium (Perissodactyla, Rhinocerotidae) from the Late Miocene of the Linxia Basin (Gansu, China). In: Cainozoic Research. 5 (1-2), 2006, pp. 93-102.
  52. ↑ Donald R. Prothero: The evolution of North American rhinoceroses. Cambridge University Press, 2005.
  53. ↑ a b Denis Geraads: Rhinocerotidae. In: L. Werdelin, DJ Sanders (Ed.): Cenozoic Mammals of Africa. Berkeley 2010, pp. 669-683.
  54. ↑ Denis Geraads, Monte MacCrossin, Brenda Benefit: A New Rhinoceros, Victoriaceros kenyensis gen. Et sp. nov., and other Perissodactyla from the Middle Miocene of Maboko, Kenya. In: Journal of Mammalian Evolution. 19 (1), 2012, pp. 57-75.
  55. ↑ Damien Becker, Pierre-Olivier Antoine, Olivier Maridet: A new genus of Rhinocerotidae (Mammalia, Perissodactyla) from the Oligocene of Europe. In: Journal of Systematic Palaeontology. 11 (8), 2013, pp. 947-972, doi: 10.1080 / 14772019.2012.699007 .
  56. ↑ Luca Pandolfi: Persiatherium rodleri, gen. Et sp. nov. (Mammalia, Rhinocerotidae) fro, the Upper Miocene of Maragheh (Northwestern Iran). In: Journal of Vertebrate Paleontology. 36 (1), 2016, p. E1040118, doi: 10.1080 / 02724634.2015.1040118 .
  57. ↑ Naoto Handa, Masato Nakatsukasa, Yutaka Kunimatsu, Hideo Nakaya: A new Elasmotheriini (Perissodactyla, Rhinocerotidae) from the upper Miocene of Samburu Hills and Nakali, northern Kenya. In: Geobios. 50 (3), 2017, pp. 197-209, doi: 10.1016 / j.geobios.2017.04.002 .
  58. ↑ Kerstin Hlawatsch, Jörg Erfurt: Tooth morphology and stratigraphic distribution of Hyrachyus minimus (Perissodactyla, Mammalia) in the Eocene Geiseltalschichten. In: Jörg Erfurt, Lutz Christian Maul (Hrsg.): 34th meeting of the working group for vertebrate paleontology of the paleontological society March 16 to March 18, 2007 in Freyburg / Unstrut. (= Hallesches Jahrbuch für Geoswissenschaften. 23). 2007, pp. 161-173.
  59. ↑ Kurt Heissig: The American genus Penetrigonias Tanner & Martin, 1976 (Mammalia: Rhinocerotidae) as a stem group elasmothere and ancestor of Menoceras Troxell, 1921. In: Zitteliana. A 52, 2012, pp. 79-95.
  60. ↑ Michael R. Voorhies, Joseph R. Thomasson: Fossil grass anthoecia within miocene rhinoceros skeletons: diet in an extinct species. In: Science. 206 (Oct 19), 1979, pp. 331-333.
  61. ↑ Pierre-Olivier Antoine, Jean Loup Welcomme: A new rhinoceros from the lower miocene of the Bugti Hills, Baluchistan, Pakistan: the earliest elasmotheriine. In: Palaeontology. 43 (5), 2000, pp. 795-816.
  62. ↑ a b Denis Geraads: Pliocene Rhinocerotidae (Mammalia) from Hadar and Dikika (Lower Awash, Ethiopia) and a revision of the origin of modern African rhinos. In: Journal of Vertebrate Paleontology. 25 (2), 2005, pp. 451-461.
  63. ↑ Deng Tao: Linxia Basin: An Ancient Paradise for Late Cenozoic Rhinoceroses in North China. In: Paleomammalogy. 24 (2), 2010, pp. 103-106.
  64. ↑ Tao Deng, Xiaoming Wang, Mikael Fortelius, Qiang Li, Yang Wang, Zhijie J. Tseng, Gary T. Takeuchi, Joel E. Saylor, Laura K. Säilä, Guangpu Xie: Out of Tibet: Pliocene Woolly Rhino Suggests High-Plateau Origin of Ice Age Megaherbivores. In: Science. 333, September 2011, pp. 1285-1288.
  65. ↑ Akira Fukuchi, Hideo Nakaya, Masanaru Takai, Shintaro Ogino: A preliminary report on the Pliocene rhinoceros from Udunga, Transbaikalia, Russia. In: Asian Paleoprimatology. 5, 2009, pp. 61-98.
  66. ↑ a b LC Rookmaaker: Early rhinoceros systematics. In: A. Wheeler et al. (Ed.): History in the service of systematics. Society for the Bibliography of Natural History, London 1981, pp. 111-118.
  67. ^ Ragnar K. Kinzelbach: An Indian Rhino, Rhinoceros unicornis, (V34) on the Artemidorus Papyros. Its position in the antique cultural tradition. In: Jochen Althöff, Sabine Föllinger, Georg Wöhrle (eds.): Ancient science and its reception. Wissenschaftlicher Verlag Trier, 2012, pp. 93-131, ISBN 978-3-86821-389-8 .
  68. ^ Wilhelm Pape and Max Sengebusch: Concise dictionary of the Greek language. 3rd edition, 6th impression, Vieweg & Sohn, Braunschweig, 1914 ( [1] )
  69. Jump up ↑ Linnaeus: Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis Editio decima, reformata. Holmiae, Laurentii Salvii, 1758, p. 56.
  70. ^ Mikael Fortelius: The morphology and paleobiological significance of the horns of Coelodonta antiquitatis (Mammalia: Rhinocerotidae). In: Journal of Vertebrate Paleontology. 3 (2), 1983, pp. 125-135.
  71. ↑ John Edward Gray: On the natural arrangement of vertebrose animals. In: London Medical Repository. 15, 1821, pp. 297–310 (306) ( PDF )
  72. ^ MC MacKenna, SK Bell: Classification of mammals above the species level. Columbia University Press, New York 1997, pp. 483-485.
  73. ↑ Richard Owen: Odontography; or, A treatise on the comparative anatomy of the teeth; their physiological relations, mode of development, and microscopic structure, in the vertebrate animals. Hippolyte Bailliere, London 1840–1845, pp. 587–599 ( [2] )
  74. ↑ Ingmar Braun, Wolfgang Zessin: Palaeolithic rhinoceros depictions and the attempt at their zoological-ethological interpretation. In: Ursus, newsletter of the Zooverein and the Schwerin Zoo. 15 (1), 2009, pp. 3-19.
  75. ↑ Michel Lorblanchet: Cave painting. A manual. Sigmaringen 1997.
  76. ^ WT Schaurte: Representation of an Elasmotherium in the rock painting of Rouffignac. In: Nature and Museum. 94 (9), 1964, pp. 354-356.
  77. ↑ Baldyrgan S. Kožamkulova: Elasmotherium sibiricum and its distribution area on the territory of the USSR. In: Quaternary Paleontology. 4, 1981, pp. 85-91.
  78. ↑ Meenakshi Dubey: Rock paintings of Pachmarhi Hills. In: Michel Lorblanchet (Ed.): Rock art in the old world. Indira Gandhi National Center for the Arts, New Delhi 2001, pp. 131-145.
  79. ↑ G. Kreuzer: Animal representations in Indian rock art. In: The Blue Booklet. 58, 1978, pp. 405-407.
  80. ↑ Sven Ouzman: Seeing is deceiving: rock art and the non-visual. In: World Archeology. 33 (2), 2001, pp. 237-256.
  81. ↑ Sven Ouzman, Jim M. Feely: Black or white? The identification and significance of rhinoceroses in South African rock art. In: Digging Stick. (SA Archaeological Society) 19 (2), 2010, pp. 9-12.
  82. ^ A. van der Geer: Rhinoceros unicornis: The Indian Rhinoceros. In: A. van der Geer: Animals in stone: Indian mammals sculptured through time. (= Handbook of Oriental Studies. Section 2: India 21). Leiden / Boston 2008, pp. 380-386.
  83. ^ WF Gowers: The classical rhinoceros. In: Antiquity. 24, 1950, pp. 61-71.
  84. ↑ HH de Longh, HHT Prins, N. van Strien, LC Rookmaaker: Some observations on the presence of one-horned rhinos in the bas reliefs of the Angkor Wat temple complex, Cambodia. In: Pachyderm. 38 (Januay-June), 2005, pp. 98-100.
  85. ^ Maryna Steyn: The Mapungubwe Gold Graves Revisited. In: The South African Archaeological Bulletin. 62 (186), 2007, pp. 140-146.
  86. ↑ Stephan Woodborne, Marc Pienaar, Sian Tiley-Nel: Dating the Mapungubwe Hill Gold. In: Journal of African Archeology. 7 (1), 2009, pp. 99-105.
  87. ↑ Alex Duffey: Mapungubwe: Interpretation of the Gold Content of the "Original Gold Burial M1, A620". In: Journal of African Archeology. 10 (2), 2012, pp. 175-187.
  88. ^ LC Rookmaaker: Captive rhinoceroses in Europe from 1500 to 1810. In: Bijdragen tot de Dierkunde. 43, 1973, pp. 39-63.
  89. ↑ International Rhino Foundation: 2007 Annual Report. Yulee, 2007.
  90. ↑ International Rhino Foundation: Annual Report 2013. ( [3] ).
  91. ↑ Richard H. Emslie, Tom Milliken, Bibhab Talukdar, Gayle Burgess, Karyn Adcock, David Balfour, Michael H. Knight: African and Asian rhinoceroses - status, conservation and trade. A report from the IUCN Species Survical Commission (IUCN / SSC) African and Asian Rhino Specialist Groups and TRAFFIC to the CITES Secretariat pursuant to Resolution Conf. 9.14 (Rev. CoP17). In: Report to CITES. 17th meeting, Colombo, CoP 18 Doc.83.1 annex 3, 2019, pp. 1–38 ( [4] ).
  92. ^ Jared Diamond: Guns, Germs, and Steel: The Fates of Human Societies. New York / London 1997, pp. 168-174, ISBN 0-393-31755-2 .
  93. ^ EB Martin: Tackling the rhino horn trade - a new approach. In: Species. 7, 1986, pp. 7-8.
  94. ^ UA Casal: Carved Rhinoceros Horns of China. In: Cultureel Indie. 2 (Aug./Sept.), 1940, pp. 212-216.
  95. ↑ a b Kyle Swenson: He smuggled a rhino horn artifact out of Miami, but the feds were watching. In: Washington Post. of October 2, 2017 ( [5] ).
  96. ↑ a b Abdul Wahab Ahmad Zafir, Junai di Payne, Azlan Mohamed, Ching Fong Lau, Dionys i us Shankar Kumar Sharma, Raymond Alfred Amirtharaj Christy Williams, Senthival Nathan, Widodo S. Ramono, Gopalasamy Reuben Clements: Now or never: what will it take to save the Sumatran rhinoceros Dicerorhinus sumatrensis from extinction? In: Fauna & Flora International, Oryx. 45 (2), 2011, pp. 225-233.
  97. ^ Mike Knight: African Rhino Specialist Group report. In: Pachyderm. 50 (July to December), 2011, pp. 7-14 ( online )
  98. ↑ RP: The first trace of the rhino mafia. ( [6] ) rp-online.de of March 3, 2012:
  99. ^ Kate Katharina Ferguson: Rhino Horn Thefts a Growing Problem in Europe. In: Spiegel Online. March 13, 2012 ( PDF ).
  100. ^ LC Rookmaaker: New literature in the Rhino Resource Center. In: Electronic Newsletter of the Rhino Resource Center. no. 25 (November), 2011, pp. 1–16 ( PDF ).
  101. ^ Felix Patton: The medicinal value of rhino horn - a quest for the truth. Unpublished report (first available on RRC), 2011, pp. 1–8 ( PDF ).
  102. ^ Mike H. Knight: African Rhino Specialist Group Chair report. In: Pachyderm. 49, 2011, pp. 6-15 ( [7] ).
  103. ^ A b c International Rhino Foundation: State of the Rhino. ( [8] ).
  104. ↑ OlPejeta Conservancy: Rhinos. ( [9] ) at: olpejetaconservancy.org.
  105. ^ International Rhino Foundation: Poaching Deterrents. ( September 10, 2014 )
  106. ^ Rhino Rescue Project: Proactive Poaching Prevention. rhinorescueproject.com from September 12, 2014.
  107. ↑ Sam Ferreira, Markus Hofmeyr, Danie Pienaar, Dave Cooper: Chemical horn infusions: a poaching deterrent or an unnecessary deception? In: Pachyderm. 55, 2014, pp. 54-61.
  108. ↑ WWF: Rhinos in Kenya: Big Brother is watching you. ( [10] ) wwf.de of March 2, 2014.
  109. ↑ Bibhab Kumar Talukdar: Asian Rhino Specialist Group report. In: Pachyderm. 49, 2011, pp. 16-19 ( (online) ).
  110. ↑ WWF: Vietnam's last rhinos should give way to the road. ( [11] ) wwf.de of May 26, 2009.
  111. ↑ WWF: Black Day for Species Protection. ( [12] ) wwf.de of October 25, 2011.
  112. ^ Susie Ellis, Maggi Moore: Conserving the Javan rhinos. In: International Zoo News. 58 (6), 2011, pp. 403-405.

Web links

Wiktionary: rhino  - explanations of meanings, word origins, synonyms, translations
Wiktionary: rhinoceros  - explanations of meanings, word origins, synonyms, translations
Commons : Rhinoceros (Rhinocerotidae)  - Collection of images, videos and audio files
Wikisource: Actual and Accurate Presentation Of The Newly Arrived Asiatic Wonder Thiers Rhinoceros  - Sources and Full Texts
This article was added to the list of excellent articles on August 5, 2012 in this version .