# Deer

Deer

Red deer ( Cervus elaphus )

Systematics
 Subclass : Higher mammals (Eutheria) Superordinate : Laurasiatheria Order : Artiodactyla (Artiodactyla) Subordination : Ruminants (ruminantia) without rank: Forehead weapon bearer (Pecora) Family : Deer
Scientific name
Cervidae
Goldfuss , 1820

The deer (Cervidae), or antlers are a mammalian family of the order Artiodactyla (Artiodactyla). The family comprises more than 80 species, of which the red deer , the fallow deer , the roe deer , the reindeer and the elk are also common in Europe . In addition, deer are found in Asia , North and South America and with a representative in Africa . The most striking characteristic of the deer are the antlers , which are varied in shape and usually only worn by the males , which are shed and newly formed every year. The main diet of the animals consists of plants, whereby soft and hard parts of plants are consumed in different amounts. Pure grazers like the hornbeams do not occur in the deer, which is related to the formation of the antlers. The social behavior of the animals is very different and ranges from solitary individuals to the formation of large, widely wandering herds. The reproductive phase is characterized by characteristic fights for dominance.

The systematic classification of deer has been the subject of numerous debates. A division into several subfamilies has often been suggested. Already in the second half of the 19th century it was recognized that the deer can be anatomically divided into two large groups, one of which is more or less restricted to Eurasia, the other to America. More recently, this dichotomy has been supported by molecular genetic analyzes. The tribal history of the deer goes back to the Lower Miocene around 20 million years ago. However, the earliest forms differed significantly from today's species and most likely did not change their forehead weapons in an annual cycle. Today's appearance and behavior emerged only in the further course of the Miocene.

## features

### General

Sika deer ( Cervus nippon ) as an inhabitant of more open landscapes
Tufted deer ( Elaphodus Cephalophus ) CLOSED senerer as residents landscapes

The size of the deer varies considerably: the length of the head and trunk varies between 70 and 310 cm, the shoulder height between 30 and 190 cm and the weight between 5.5 and 770 kg. The smallest living representative is the southern pudu ( Pudu puda ), the largest of the elk ( Alces alces ). Most species, with the exception of the muntjak ( Muntiacus muntjak ), have a sexual dimorphism in terms of size. In the Tenasserim muntjak ( Muntiacus feae ) or in the water deer ( Hydropotes inermis ) the females are mostly larger than the males, with all other representatives the male is significantly larger and heavier than the female. The body shape is also variable; within the family, two general construction plans can be distinguished. One includes animals with a stocky build, short neck, rounded back, and strong hind legs and less well developed front legs. These are mostly smaller representatives such as the pudus , the muntjak deer or the spit deer , which are still reminiscent of primitive artifacts and have only short, pike- like antlers. They are often inhabitants of dense forests or of landscapes with lush vegetation, in which they can jump around quickly. In addition, the second building plan is characterized by slender animals with comparatively long limbs and multiple articulated antlers. The relatives mostly live in more open landscapes and are good runners. In both construction plans the tail is rather short like a stub. The fur predominantly having in the forest residents a camouflaged brown or gray color, more open with some types landscapes such as Prince Alfred Hirsch ( Rusa alfredi ) or the axis deer ( Axis axis ), a dotted fur apparel has been formed, otherwise accents often occur of the head and the rear part. The head is usually elongated, the ears are large and erect. In addition, deer have three types of glands : Almost all representatives have pre-eye glands , and apart from the Muntjak deer, metatarsal brushes are found on the hind legs. Most representatives of the deer also have interdigital glands .

### antler

Different types of deer antlers

Characteristic for the deer is the antlers , a pair of formation that grows from cone-shaped bones ("rose bushes") on the frontal bone (os frontale). The antlers consist of bone substance, the largest part is made up of hydroxyapatite , a crystallized calcium phosphate that makes up a good 30% of the antlers. The shape of the antlers depends on the age and the species, in some species it is simple, pike-shaped, in others it has widely branched or shovel-shaped structures with numerous tips. The elk has the largest antlers of today's deer, weighing up to 35 kg. The large and widely ramified antlers often have a cancellous core, which increases the elasticity of the entire structure. With the exception of the water deer, all deer species have antlers. However, this is only developed in both sexes in the reindeer ( Rangifer tarandus ), otherwise it is an exclusive feature of the males. In these, it serves to showcase and fight for the mating privilege.

In contrast to the horns of the hornbeams, the antlers are not a permanent formation, but are renewed in an annual cycle. The formation is linked to the testosterone balance of the males (in the case of Ren possibly also to the estradiol , which is produced in larger quantities in both sexes). During the growth phase, the antlers are supplied with blood through a short-haired skin called bast. The growth rate is enormous and can for large dogs such as the elk ( Cervus canadensis are cm daily) in 2.8 g or be in elk 417, whereby the antlers one of the fastest growing organs showing inside of mammals. However, the formation of the antlers is very costly, as minerals from the skeleton must be mobilized in some cases, which in some species leads to temporary osteoporosis . Once it reaches its full size, the skin's outer layer dries up and becomes itchy from the blood supply to dry, which is why it is peeled off or swept away . What remains is a dark and largely dead bone structure that is connected to the living "rose bush" and is worn for several months.

Different growth stages of an antler

The antlers are shed every year after the mating season and then re-formed. The shedding of the antlers is associated with a drop in testosterone levels, which briefly activates osteoclasts that dissolve the bones on the "rose bush". The subsequent healing of the wound is probably the trigger for the next antler growth. With most deer, there is a time lag of one to two months between shedding and new formation, with real deer this can follow one another. In species with a fixed mating season, this shedding occurs in a certain season (in the case of deer and male reindeer in late autumn, in female reindeer and other European species in late winter or spring); for species in tropical regions there is no fixed point in time for this. The formation of the antlers as early as the juvenile a stage and begins with the illusion deer and Muntjakhirschen in the first year, with the Real deer for the second year. Smaller skewers are formed first, the complex antler structures arise with increasing age.

Development of the deer antlers

According to phylogenetic analyzes, the last common ancestor of today's deer had two-part antlers with the (lower) antler rod and the eye shoot. This original variant is only preserved today with the Muntjak deer. Three-pointed antlers developed independently within the real deer and the deer. There was also a lot of redesign. Within the real deer, the complex antlers of the red deer ( Cervus elaphus ) and the elaphus , which ideally consist of the eye shoot, the ice shoot, the central shoot and the crown with three ends, which are connected to each other via the lower and higher pole section, also emerged . This can be found almost identically in the sika deer ( Cervus nippon ), which, however, lost its ice sprout. The fallow deer ( Dama dama ) also formed a lead sprout and a shovel-shaped widening of the higher pole section. The Barasinghas, on the other hand, reduced the ice rungs and the higher pole section, but developed a vertical and a rear pole section with additional rungs. In the case of the lyre deer and the Davidic deer ( Elaphurus davidianus ), the higher pole section is largely absent, but there are also several cacuminal rungs and a medial branch on the lower pole section, as in the latter species the eye branch enlarged and divided into two ends. The antlers developed somewhat differently within the deer , the most original form of which can still be found in the roe deer ( Capreolus capreolus ). The complex, higher pole section is missing, but a forehead sprout has separated. Other forms instead of the higher one have a secondary upper pole section with additional rungs such as the back sprout and the end rungs, for example in the case of the reindeer or the representatives of the real deer . The elk, on the other hand, does not have an eye or frontal shoot, but rather oversized end shoots. Only the water deer lost all of its antlers.

### Skeletal features

Skeleton of the water deer (
Hydropotes inermis )
Giant Muntjac Skull (
Muntiacus vuquangensis )

The skull generally has a long, narrow construction with an elongated rostrum . The nasolacrimal duct (ductus nasolacrimalis) is bifurcated and there are two tear holes (foramina lacrimalia) at the front edge of the eye socket (orbit). The upper incisors are always missing, in the lower jaw there are three per jaw half. The upper canine tooth is enlarged in species with missing or small antlers ( water deer , muntjacs ) and protrudes like a tusk from the mouth, in the other species it is reduced in size or completely absent. The lower canine resembles the incisors and forms a closed row with them. There are three premolars and three molars per half of the jaw , which are rather low-crowned ( brachyodont ). A crescent-shaped, longitudinal enamel pattern ( selenodont ) is formed on the chewing surface . Overall, the following tooth formula results : That means 32 to 34 teeth are formed. The lower jaw is very strong and has a wide angular process as an attachment point for the masticatory muscles. ${\ displaystyle {\ frac {0.0 (1) .3.3} {3.1.3.3}}}$

Anatomical training of the front legs in "Telemetacarpalia" (left) and "Plesiometacarpalia" (right)

As with all cloven-hoofed animals , the central axis of the foot lies between rays III and IV, which are enlarged and the only ones touching the ground, so the animals stand on toes 3 and 4. The corresponding third and fourth metacarpal and metatarsal bones are like that called cannon leg fused. The first toe is completely missing, the second and fifth toes are greatly reduced in size and no longer touch the ground. The degree of reduction of the second and fifth toes forefoot an important criterion to distinguish the two major directions: Cervinae ( cervinae and muntiacini ) are "Plesiometacarpalia" (from greek πλησίον ( plesion "near") = and μετακάρπιον ( metakarpion ) = " Metacarpal "), which means that the proximal (near the center of the body) parts of the 2nd and 5th metacarpals (metacarpal bones) are present, the respective three phalanges are also formed, but are separated from the metacarpal bones by a large gap. On the other hand represent the capreolinae (Capreolinae; Actual capreolinae , deer and elk) "Telemetacarpalia" (from the Greek τήλε ( tele ) = "far" and μετακάρπιον ( metakarpion ) = "median") represents, that is, that only the distal (from The metacarpals are formed, the phalanxes associated with them articulate directly with the metacarpal bones. The shape and spreadability of the hooves depend on the inhabited landscape and range from further apart hooves in residents of damp to swampy biotopes to wide hooves in those in snowy areas.

## distribution and habitat

Roe deer ( Capreolus capreolus ), the most common species of deer in Central Europe

The natural range of the deer includes large parts of Eurasia and America , they reach their highest diversity in South America and Southeast Asia . In Africa they are only found in the northwestern part, in the areas south of the Sahara they are absent and are replaced there by the hornbeams . Deer have been introduced by humans to some regions where they were not native, including Australia , New Zealand , New Guinea, and some Caribbean islands . Deer inhabit a wide variety of habitats. The majority of the species prefer closed forests, more open forest landscapes and forest edges. Some forms, such as the Muntjak deer or the spit deer , are generally to be found in dense vegetation, while others show a more plastic behavior and mostly occur in forest edge locations with transition to more open landscapes. Few representatives such as the reindeer or the Pampas deer ( Ozotoceros bezoarticus ) fit directly in open country, in addition to live among others, the marsh deer ( Blastocerus dichotomus ) and the water deer in swamp - or marshes . The deer have opened up both lowlands and highland locations up to 5100 m and inhabit tropical climates as well as the arctic tundra .

## Way of life

### Territorial and social behavior

Reindeer ( Rangifer tarandus ) in herd

Deer are faithful to their location or migrate regularly. Species that live strictly in the forest, such as the pudus, mazamas or the crested deer, are solitary and mostly live hidden in the forest thicket. They escape predators by fleeing quickly with wide jumps, which their often stronger hind legs make possible. The animals live in action spaces that can be between 15 and 20 ha for the southern pudu ( Pudu puda ), up to 100 ha for the great mazama ( Mazama americana ) or the Chinese muntjak ( Muntiacus reevesi ). The territories are partially defended territorially, secretions from the glands on the head and feet and also urine are used to communicate with conspecifics and to mark the boundaries of the territories, and sound communication is usually minimized. Territoriality can also be restricted in poor food conditions and is then reduced to dominance. Pair formations only occur during the mating season or are found in mother-young groups. In some species there is also only a temporary territoriality that is limited to the reproductive phase, such as the roe deer ( Capreolus capreolus ).

Axis deer ( Axis axis ) in the family

However, the majority of species live in groups, the size of which can vary according to species and habitat. In group formations, both herds of females with their young and bachelor groups of young adult males occur, mixed associations usually only arise during the mating season. The hikes are mainly triggered by the search for the best food sources. In the more tropical regions, the hikes are generally short and mostly take place in open landscapes, where the animals look for fresh food during the dry season or leave floodplain areas. In mountain regions there is often a seasonal change in altitude, which usually only covers a few kilometers. In addition, there are also real migrations , especially in the temperate and cool climate zones , in which long distances are covered. The Siberian deer ( Capreolus pygargus ), for example, migrates up to 500 km over the course of the year. The most spectacular are those of the reindeer ( Rangifer taranduss ), which form herds of up to 500,000 individuals and can roam around 5000 km in the tundra regions within a year, covering 20 to 50 km daily. The extent to which animals are sedentary or wander around depends on the external conditions. Reindeer in forest regions migrate less often than those on the tundra. The elk ( Cervus canadensis ) has both local and migratory animals. Hikers are exposed to a higher mortality , but often open up better grazing grounds.

### nutrition

David deer ( Elaphurus davidianus )

Deer are generally herbivores , and food consumption takes up a large part of the daily workload. The animals eat in phases, with a day having between five and eleven eating phases depending on the size of the stomach. In between there are long periods of rest during which the food is digested and chewed on again. The animals feed on different parts of plants such as leaves , bark , buds and twigs , but also on fruits and, more rarely, on grasses . They are very adaptable, but compared to the horn-carriers they generally prefer a softer plant food, which is also indicated by the lower average tooth crowns and thus more brachyodontic teeth. As a result, there are no real grazers within the deer as in the hornbills, which means that no representative is adapted to extremely dry landscapes. A purely grass-eating diet is also incompatible with the formation of antlers, since grasses are very low in energy and contain too few minerals, which are essential for the development of the larger and heavier antlers in particular. The inhabitants of closed forests with still relatively primitive physiques such as the muntjaks, pudus and deer feed mainly on leaves ( browser ), some of the Mazama species are fruit-eaters, as has also been partially proven for the earliest forms of deer and their ancestors. The proportion of fruit can exceed two thirds of the amount of food consumed. The leaf eaters tend to be opportunistic and eat a wide variety of plants, but usually only the most easily digestible parts of the plant. On the other hand there are the group deer, the reindeer or some noble deer such as the elk and the white-lipped deer, which tend to a grass-eating way of life ( grazer ). However, to a certain extent they also eat softer foods such as aquatic plants , herbs or lichens . The majority of the species eats a mixed plant food ( mixed feeder ) and can thus eat soft and hard plants depending on the circumstances and sometimes seasonal availability.

Numerous representatives also eat animal food, such as crustaceans , birds or fish . Furthermore, the gnawing of bones or antlers is known, which can often be observed in male animals. The behavior helps to balance or increase the mineral balance during antler growth and can also alleviate or suppress the osteoporosis that is sometimes associated with it . Species that are particularly widespread in temperate and cold climates have an annual cycle in their food intake, which includes different amounts and compositions. This annual cycle begins in spring for females with the end of the gestation period and the onset of milk production for the offspring, which requires large amounts of food. During this time, males eat a high proportion of fat, which they consume in autumn during the rutting phase , during which they hardly eat any food. In winter the animals generally consume a smaller amount of food, which is associated with a decrease in the metabolic rate and saves energy for maintaining the body temperature.

### Reproduction

European elk ( Alces alces ), cow with young animal

The mating times of the individual deer species, commonly referred to as rut , vary depending on the geographical region. In tropical landscapes, reproduction can take place all year round, in temperate regions it usually takes place in autumn or winter and is therefore seasonal. The sexual cycle varies from 11 days in the pudus to 29 days in the axis deer. The female animals are usually only ready to receive for a short time, which lasts about 12 to 24 hours. As mostly polyestrial animals , ovulation takes place cyclically until fertilization occurs. An exception is the deer , where mating takes place in summer (July / August), during which the females are ready to receive for a maximum of 36 hours; in doing so, they often undertake rambling hikes far from their areas of action in order to increase the possibility of mating. The gestation period is when the deer usually six to nine months, here is the deer is an exception, as through a dormancy discharging the offspring will be extended to ten months. The litter size is usually one boy, only two to a maximum of five kittens are born in the deer. These are refugees and usually wear a spotted coat of fur. In almost all species of deer, the boys hide in thickets initially ( Ablieger ). Rearing is mostly done by the mother.

Fighting males of white-tailed deer ( Odocoileus virginianus )

The males of smaller species are rarely polygynous . During the reproductive phase, they form temporary pairs with one or two females who only stay together for two or three days. There are different reproductive strategies for strongly polygynous species. The males penetrate the territories of the females and draw attention to themselves in the form of markings with scented secretions or strong vocalizations with repeated loud rutting cries. Others form harems and defend them against competitors (" top dog "), lead dominance fights in mixed groups or fight for mating privilege in courtship areas . Different strategies can occur within a species, for example with fallow deer or red deer, which adapt their behavior to the given situation and are able to vary their pattern within a day. The fights of the males are carried out with the tusk-like canine teeth or the antlers. They are ritualized and follow a given choreography, which in antlered animals involves lowering the head, then hooking the antlers and circling each other. In the sambar, collision fights similar to the ibex are known. As a rule, the animals avoid serious injuries, but these can certainly occur. In strongly dimorphic species, the mating right is mostly reserved for the larger and stronger males due to the fighting, with fallow deer 3% of the males cover an average of three quarters of the receptive females.

## Systematics

### External system

Internal systematics of the recent cetartiodactyla according to Zurano et al. 2019
Cetartiodactyla
Suina (pig-like)
 Tayassuidae (umbilical pigs) Suidae (real pigs)

Camelidae (camels)

Cetancodonta
 Hippopotamidae (hippos) Cetacea (whale)

Ruminantia (ruminant)

Tragulidae (deer piglets)

Giraffidae (giraffes)

Cervidae (deer)

 Moschidae (musk deer) Bovidae (hornbeam)

The deer are one of the six surviving families of the subordination of the ruminants (ruminant) within the order of cloven-hoofed animals (Artiodactyla). Within the ruminant turn the deer together with the horn carriers (Bovidae), the giraffe-like (Giraffidae), the musk deer (Moschidae) and the Gabelhorn carriers (Antilocapridae) the parent taxon the end weapon carrier (Pecora) to which the mouse deer (Tragulidae) as a sister group against stand. For a long time the musk deer were considered to be closely related to the deer and were sometimes integrated into them as a subfamily. From a molecular genetic point of view, however, they form an independent group, although it is still unclear whether they are more closely related to the horned bearers or to the deer. The former variant currently seems the more likely. In some cases, the forked horn carriers with the forkbuck ( Antilocapra ) were considered to be very close to the deer due to the structure of the lacrimal bone and were combined with these to form the superfamily of the cervoidea (deer-like). However, this contradicts the molecular genetic investigations, which classify the fork horn bearers more at the base of the forehead weapon bearers.

### Internal system

The common feature of all deer is the absence of the gallbladder and the formation of antlers, which are renewed annually. Today they are divided into two subfamilies with a total of five branches, which differ among other things in the arrangement of the reduced toes (see above), in the antlers and in the details of the skull structure. The subfamily of the Cervinae includes the Cervini (real deer) and the Muntiacini (Muntjak deer) , both groups are characterized by the "plesiometacarpale" toe position. The second subfamily, the Capreolinae , consists of the Capreolini , the Alceini, the Rangiferini and the Odocoileini (actual deer ) and corresponds to the "Telemetacarpalia" group. The alceini and the odocoileini, in turn, are united by the presence of interdigital glands ( glands between the toes), while the odocoileini, often also referred to as neocervines , are characterized by the fact that the ploughshare is completely integrated into the nasal septum and thus the choane , the posterior section of the Nostril, vertically divides. Traditionally, the antlerless water deer ( Hydropotes inermis ) was compared to the other deer as the most primitive representative. Molecular genetic studies, however, reveal a close relationship with the deer ( Capreolus ); a separation from the deer would make them appear paraphyletic . Accordingly, the lack of antlers in water deer may not be an original, but a derived feature. The same applies to the Muntjak deer, in which the genetic data suggest a close relationship with the real deer. The division of the deer into the two subfamilies, according to molecular genetic studies, did not take place until the Upper Miocene, around 9 million years ago, a rapid radiation of the individual lines can therefore be assumed for the end of the Miocene and the Pliocene .

### Overview of the genera and species of deer

Internal systematics of the Cervidae

according to Heckeberg 2020

(mtDNA and core DNA combined)

Cervidae

Internal systematics of the cervinae

according to Heckeberg 2020

(mtDNA and core DNA combined)

Cervinae
Cervini

 Megaloceros (†)

Muntiacini

Internal systematics of the Capreolinae according to Heckeberg 2020

(mtDNA and core DNA combined)

Capreolinae

Odocoileini
Blastocerina

 Ozotoceros (including "Hippocamelus" antisensis )

Mazama ( M. gouazoubira , M. chunyi )

Odocoileina

Mazama ( M. americana , M. bororo , M. nana , M. temana )

Mazama ( M. americana , M. rufina , M. bricenii )

Rangiferini

Alceini

Capreolini

The following differences occur in a purely mitochondrial cladogram:

• the position of the elk is basal within the Capreolinae
• the position of the north pudu is basal within the blastocerina.

The following overview is largely based on the work of Colin Peter Groves and Peter Grubb 2011 and on S. Mattioli 2011. The classification above the genus level was made in 2006 by Gilbert et al. suggested:

• Family Cervidae Goldfuss , 1820
• Tribus Muntiacini Knottnerus-Meyer , 1907 ( Muntjak deer )
• Genus Elaphodus Milne-Edwards , 1872
• Tufted deer ( Elaphodus Cephalophus Milne-Edwards , 1872)
• Genus Muntjaks ( Muntiacus rafinesque , 1815)
• Tribus Cervini Goldfuss , 1820 ( real deer )
• Genus Axishirsche or Spotted Deer ( Axis C. H. Smith , 1827)
• Genus Elaphurus Milne-Edwards , 1866
• Genus Barasinghas or grouper deer ( Rucervus Hodgson , 1838)
• Manipur lyre deer ( Panolia eldii ( McClelland , 1842); also Rucervus eldii )
• Thailand lyre deer or Siamese lyre deer ( Panolia siamensis ( Lydekker , 1915); also Rucervus siamensis )
• Myanmar lyre deer or Burma lyre deer ( Panolia thamin ( Thomas , 1918); also Rucervus thamin )
• Genus red deer or red deer ( Cervus Linnaeus , 1758)
• Subfamily Capreolinae Brookes , 1828 (often Odocoileinae capreolinae )
• Genus Hydropotes Swinhoe , 1870
• Water deer or Chinese water deer ( Hydropotes inermis Swinhoe , 1870)
• Genus deer ( Capreolus gray , 1821)
• Tribus Alceini Brookes , 1828
• Genus Elche ( Alces Gray , 1821)
• European elk or Eurasian elk ( Alces alces ( Linnaeus , 1758))
• American elk or East American elk ( Alces americanus ( Clinton , 1822))
• Tribus Rangiferini Brookes , 1828
• Genus Rangifer C. H. Smith , 1827
• Ren and reindeer ( Rangifer tarandus ( Linnaeus , 1758))
• American deer genus ( Odocoileus Rafinesque , 1832)
• Genus Blastocerus Wagner , 1844
• Genus Ozotoceros Ameghino , 1891
• Genus Pudus ( Pudu Gray , 1852)

The systematics of the Neotropical Deer (Odocoileini) is currently very problematic and will most likely have to be rearranged in the near future. The pike deer ( Mazama ) are paraphyletic according to molecular genetic studies and are divided into two different clades, which include the Andean deer, the marsh deer and the pampas deer on the one hand, and the American deer on the other. As a result, the genus of the common deer has to be split into several groups. Within the clade with the American deer, a union with Mazama is possible due to the close genetic relationship , on the one hand a merger of Odocoileus with Mazama , on the other hand only with "Mazama" pandora was proposed . The problem is that the American deer obviously do not form a closed group either, as some of the subspecies of the mule deer ( Odocoileus hemionus ) group more closely with “Mazama” pandora , while another group with the white-tailed deer ( Odocoileus virginianus ). As a result of further analyzes, the Great Mazama ( Mazama americana ) is to be regarded as a species complex . Likewise, the pudú (set Pudu ) probably not represent a common group, as the Südpudu is closer to the sump Hirsch and Pampa Hirsch, the Northern Pudu contrast, the Schwestertaxon represents all other authentics illusion deer and reindeer, for the latter a displacement in the genus was Pudella proposed. The Andean deer ( Hippocamelus ), which contain two highly variable species, may also be regarded as paraphyletic. Within the cervinae, the pink deer must be classified as paraphyletic. One group is associated with the Axis deer, the other is closer to the red deer. The latter includes the lyre deer and was therefore moved to the genus Panolia . The white-lipped deer ( Cervus albirostris ) was originally assigned to the genus Przewalskium , while the sambar deer formed the genus Rusa . However, genetic studies show that Przewalskium and Rusa are deeply embedded in the genus Cervus . Alternatively, it was proposed to also incorporate the lyre deer and the David deer ( Elaphurus davidianus ) in Cervus . A stronger subdivision of the red deer itself was proposed for the eastern species group (elk and sika deer) as early as 2006.

## Tribal history

Skull of Procervulus

The origin of the deer is believed to be in Eurasia. The Dromomerycidae , which occurred in the Lower and Middle Miocene in North America and were characterized by unusually shaped forehead weapons, are very closely related to the deer . The earliest deer appeared around the same time, around 20 million years ago, but their evolutionary history is still largely unclear. The early forms are mostly assigned to their own subfamilies such as the Lagomerycinae or the Procervulinae . They also wore antler-like structures that differed significantly from those of today's deer. The antlers were relatively small, consisted of long cones and branched out only shortly before the end. They rose up steeply and, similar to today's representatives, often only occurred in males. It is possible that the forehead weapons were not discarded annually due to the lack of the "rose bush", but existed permanently or only the uppermost, branched part was subject to an annual renewal cycle. In terms of skeletal anatomy, the early deer resembled the musk deer (Moschidae), with the molars already largely corresponding to the deer, while a large, backward-curved canine tooth appeared. The oldest forms include Lagomeryx , Ligeromeryx or Procervulus . In general, a leafy to fruit-based diet is assumed for the early deer, but studies on the teeth of Procervulus suggest that at least in this case, a certain proportion of grass also played a role.

Skeleton of Megaloceros

In the course of the Middle and Upper Miocene, the front cones shortened, the upper part of the antlers lengthened and "rose bushes" developed. In addition, the antlers moved a little backwards on the skull and the horn cones no longer protruded so steeply. The dentition also went through some changes with more strongly molarized premolars and higher-crowned posterior molars. In addition, the animals increased significantly in body size. In the late Middle Miocene 11 million years ago, the first deer appeared, which were already very similar to today's species. With Euprox , a first representative of the Muntjak deer from this period has possibly been proven, early finds come from Central Europe and East Asia. In the late Miocene, about 9 to 7 million years ago, finds from southeastern China date back to Muntiacus and thus to a form that still occurs today. At about the same time, Procapreolus as a relative of the roe deer and Cervavitus as the forerunner of the elk are proven. The first large-shaped representatives of the Megacerotini (also Megalocerotini) appear for the first time in this period, which are mainly known from the Pleistocene for their shapes with strikingly large antlers, including Eucladoceros and the "giant deer" Megaloceros - the size of a large elk was comparable, but had significantly wider antlers than all deer species living today.

The earliest record of deer in North America comes from the Lower Pliocene around 5 million years ago with Eocoileus from the palmetto fauna of Florida. They only reached South America when the land connection of the Isthmus of Panama closed 3 million years ago . The first immigrants were mostly medium-sized animals with multi-forked antlers such as Antifer , which appeared in western South America around 2 million years ago.

## Research history

August Goldfoot

The systematic classification of the deer has a controversial past and has not yet been fully clarified. The family is named after August Goldfuß , who coined the term “Cervina” (from Latin cervus “deer”) in 1820 and united the “deer animals ” under it. According to Goldfuß, these included the genus Cervus , to which he assigned not only the red deer, but also the roe deer, the fallow deer, the reindeer and the elk, as well as the giraffes and the musk deer . Only a year later, John Edward Gray first used the name Cervidae, which is in use today, within which he differentiated Cervus , Alces and Capreolus . Together with other cloven-hoofed animals, he referred the deer to the group of ruminantia (ruminants). More than 50 years later, in 1872, Gray had already split the deer into four families in a catalog of the British Museum , into the Cervidae (deer), Alceidae (elk), Rangiferidae (ren) and Cervulidae (Muntjak deer). At that time he referred the water deer to the musk deer (Moschidae). In an important attempt at classifying deer by Victor Brooke in 1878, he differentiated the family into the "Plesiometacarpalia" and the "Telemetacarpalia", using the formation of the second and fifth toes of the forefoot as a criterion. Brooke recognized that the fundamental division of the deer into two parts was also repeated in the skull features, for example in the formation of the ploughshare . In addition, it was also geographically, as the "plesiometacarpalia" are mainly restricted to Eurasia (with the exception of the North American elk), but the "telemetacarpalia" mostly only occur in America (with the exception of the Eurasian deer as well as the elk and the reindeer, which are circumpolar occur).

Later authors took Brooke's findings into account, the structure of the deer remained controversial, but in many cases reflected Brooke's central division. Reginald Innes Pocock identified a total of eight subfamilies in 1923, which he determined mainly on the basis of skull features. In his general classification of mammals, George Gaylord Simpson divided today's deer into three subfamilies (Muntiacinae, Cervinae and Odocoloinae) in 1945, but also placed the musk deer as a fourth. He also split the three subfamilies of the deer into six tribes. Overall, the system was very similar to that of Brooke, the Muntjak deer Simpson saw in its structure as the most primitive group. Konstantin K. Flerov in turn criticized Simpson's system of deer in 1952 and introduced another subfamily for water deer, the Hydropotinae. As early as 1974 Colin Peter Groves pointed out that the Muntjak deer also have a "plesiometacarpals" forefoot and therefore, in addition to some other skull features, can be combined with the cervinae. The conception was later adapted by Don E. Wilson and DeeAnn M. Reeder for their 2005 work Mammal Species of the World , in which they highlighted three subfamilies with the Cervinae, the Capreolinae and the Hydropotinae.

The molecular genetic analysis methods that emerged towards the end of the 20th century also revealed that deer were divided into two parts. This showed that the Muntjak deer are actually closely related to the real deer, whereas the water deer has a close connection to the deer. Consequently, only two subfamilies of deer are recognized today, which largely correspond to the classic subdivision according to Victor Brooke.

## Deer and people

### designation

The word deer goes back via Old High German hir (u) z as in Latin cervus to the Indo-European roots kerud- (horned, consecrated, stag) and * ker , which means “horn” or “antler”. In this respect, the male animal is the deer , not wrongly the bull . The naming of the sexes is biologically ambiguous. In species whose name ends in -hirsch , females are often called hind ; outdated names for this are hind or hindin . Young animals are called deer calves . In the case of the deer species known as deer , however, males are called buck ; Females are called doe or goat and young animals are called fawns . For reindeer, on the other hand, the Sami names Sarves are sometimes used for the male and Vaia for the female.

### Benefits and threats

Deer have been hunted by humans for various reasons since the early days. On the one hand, this includes the benefits of eating their meat and processing their fur. On the other hand, it is often a hunt for purely sporting reasons, which mainly affects the males because of their antlers. One species, the reindeer , was domesticated and not only serves as a supplier of meat and fur, but is also kept as a milk and pack animal. Several species were introduced to other regions as hunting and park animals, for example fallow deer can be found in Northern and Central Europe, water deer in Western Europe and red deer in Australia and New Zealand.

This increase in the range of some species is offset by the threat from a number of species, which lies in hunting on the one hand and in the destruction of their habitat on the other. One species, the Schomburgk deer , became extinct in the 20th century, a second species, the David deer , only escaped this fate through breeding in European enclosures, but is still considered critically endangered. The IUCN lists four types as endangered ( endangered ) and six species at risk ( endangered ), for some species, however, lack accurate data.

### Deer in culture

Statue of the goddess Artemis with a deer

Deer also found their way into mythology and culture in various peoples. Already in Paleolithic rock carvings, both animals and people are depicted with deer antlers. In Celtic mythology , Cernunnos is an embodiment of the "deer god", a type that can also be found in other religions. In ancient Greece, the stag was sacred to Artemis, the goddess of hunting . Agamemnon killed such a doe in Aulis , whereupon Artemis sent a calm and demanded Agamemnon's eldest daughter Iphigenia as an atonement . Another sacred doe was the Kerynite doe , who had golden antlers and lived in Arcadia. Heracles had to catch them alive as a third task. The Norse mythology knows deer, so eat four of these animals , the buds of the world ash Yggdrasil .

In the Christian context, the stag is found as a result of Psalm verse 42.2 ("As the stag thirsts for fresh water, so my soul, God, thirsts for you.") As a representation of the soul seeking salvation or saved in baptism and is therefore also often depicted on baptismal fonts and in baptisteries . Legends of saints tell of encounters with deer with a crucifix in their antlers, for example with the Saints Eustachius or Hubertus von Liège . In the Bible, the deer is also often a symbol of speed and agility, as in Psalm 18:34 ("He made me leap like deer") or in Isaiah 35: 6 ("Then the lame leaps like a deer"). With reference to Psalm 42: 2 and the Physiologus , enmity towards the serpent or the dragon has been an essential characteristic of the deer since ancient times. Thus, in its role as an opponent of the snake, i.e. the devil, the stag has a christological significance, even if it appears less often as a representative of Christ than the fish or the lamb in Christian symbolism . According to the Christian tradition, the deer counts as a particularly clean animal, since it feeds exclusively on plants, which is why its meat is considered to be cleansing. Also Hildegard von Bingen characterized in their Physica the deer as an enemy of " toad " and refers to the healing and cleansing effect of deer meat . Due to the annual shedding and redevelopment of its antlers, the deer is also a symbol of resurrection and renewal.

Numerous medieval texts tell of wonderful deer hunts, of stags that lead the hero into fairyland to win the love of a fairy, but also of stags that show the deceased the way to the realm of the dead.

In the non-European area, deer appear in Shinto , where they are considered divine messengers and are kept, for example, around the Kasuga shrine . Deer - primarily elk - also play a role in Indian mythology and are associated with qualities such as gentleness and gratitude.

## literature

• Don E. Wilson, DeeAnn M. Reeder (Eds.): Mammal Species of the World. 3rd edition. Johns Hopkins University Press, Baltimore 2005, ISBN 0-8018-8221-4 .
• Hubert Hendrichs, Roland Frey: Cervidae, deer. In: Wilfried Westheide, Reinhard Rieger (Ed.): Special Zoology. Part 2: vertebrates or skulls. Spektrum Akademischer Verlag, Heidelberg / Berlin 2004, ISBN 3-8274-0307-3 , pp. 628–630.
• S. Mattioli: Family Cervidae (Deer). In: Don E. Wilson, Russell A. Mittermeier (eds.): Handbook of the Mammals of the World. Volume 2: Hooved Mammals. Lynx Edicions, Barcelona 2011, ISBN 978-84-96553-77-4 , pp. 350-443.
• Ronald M. Nowak: Walker's Mammals of the World. Johns Hopkins University Press, Baltimore 1999, ISBN 0-8018-5789-9 .

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