Antler

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Mature red deer stag, Denmark (2009) Red deer stag 2009 denmark.jpg
Mature red deer stag, Denmark (2009)
Red deer at the beginning of the growing season Antlers at the start of the season, cervus elaphus, deinze, belgium.jpg
Red deer at the beginning of the growing season

Antlers are extensions of an animal's skull found in members of the Cervidae (deer) family. Antlers are a single structure composed of bone, cartilage, fibrous tissue, skin, nerves, and blood vessels. They are generally found only on males, with the exception of reindeer/caribou. [1] Antlers are shed and regrown each year and function primarily as objects of sexual attraction and as weapons in fights between males for control of harems.

Contents

In contrast to antlers, horns—found on pronghorns and bovids, such as sheep, goats, bison and cattle—are two-part structures that usually do not shed. A horn's interior of bone is covered by an exterior sheath made of keratin [2] (the same material as human fingernails and toenails).

Etymology

Antler comes from the Old French antoillier (see present French : "Andouiller", from ant-, meaning before, oeil, meaning eye and -ier, a suffix indicating an action or state of being) [3] [4] possibly from some form of an unattested Latin word *anteocularis, "before the eye" [5] (and applied to the word for "branch" [6] or "horn" [4] ).

Structure and development

Male fallow deer fighting KaempfendeHirsche-2-cropped.jpg
Male fallow deer fighting
Two Sambar deer fighting, Silvassa, India Sambar deers Fighting Silvassa.jpg
Two Sambar deer fighting, Silvassa, India

Antlers are unique to cervids. The ancestors of deer had tusks (long upper canine teeth). In most species, antlers appear to replace tusks. However, one modern species (the water deer) has tusks and no antlers and the muntjac has small antlers and tusks. The musk deer, which are not true cervids, also bear tusks in place of antlers. [7]

Antlers are usually found only on males. Only reindeer (known as caribou in North America) have antlers on the females, and these are normally smaller than those of the males. Nevertheless, fertile does from other species of deer have the capacity to produce antlers on occasion, usually due to increased testosterone levels. [8] The "horns" of a pronghorn (which is not a cervid but a giraffoid) meet some of the criteria of antlers, but are not considered true antlers because they contain keratin. [9]

An antler on a red deer stag. Velvet covers a growing antler, providing blood flow that supplies oxygen and nutrients. Red deer stag velvet.jpg
An antler on a red deer stag. Velvet covers a growing antler, providing blood flow that supplies oxygen and nutrients.

Each antler grows from an attachment point on the skull called a pedicle. While an antler is growing, it is covered with highly vascular skin called velvet, which supplies oxygen and nutrients to the growing bone. [7] Antlers are considered one of the most exaggerated cases of male secondary sexual traits in the animal kingdom, [10] and grow faster than any other mammal bone. [11] Growth occurs at the tip, and is initially cartilage, which is later replaced by bone tissue. Once the antler has achieved its full size, the velvet is lost and the antler's bone dies. This dead bone structure is the mature antler. In most cases, the bone at the base is destroyed by osteoclasts and the antlers fall off at some point. [7] As a result of their fast growth rate, antlers are considered a handicap since there is an immense nutritional demand on deer to re-grow antlers annually, and thus can be honest signals of metabolic efficiency and food gathering capability. [12]

Increasing size of antlers year on year in different European game species, 1891 illustration Geweihe Pierer.jpg
Increasing size of antlers year on year in different European game species, 1891 illustration

In most arctic and temperate-zone species, antler growth and shedding is annual, and is controlled by the length of daylight. Although the antlers are regrown each year, their size varies with the age of the animal in many species, increasing annually over several years before reaching maximum size. In tropical species, antlers may be shed at any time of year, and in some species such as the sambar, antlers are shed at different times in the year depending on multiple factors. Some equatorial deer never shed their antlers. Antlers function as weapons in combats between males, which sometimes cause serious wounds, and as dominance and sexual displays. [11]

Function

Sexual selection

The principal means of evolution of antlers is sexual selection, which operates via two mechanisms: male-to-male competition (behaviorally, physiologically) and female mate choice. [10] Male-male competition can take place in two forms. First, they can compete behaviorally where males use their antlers as weapons to compete for access to mates; second, they can compete physiologically where males present their antlers to display their strength and fertility competitiveness to compete for access to mates. [10] Males with the largest antlers are more likely to obtain mates and achieve the highest fertilization success due to their competitiveness, dominance and high phenotypic quality. [10] Whether this is a result of male-male fighting or display, or of female choosiness differs depending on the species as the shape, size, and function of antlers vary between species. [13]

Heritability and reproductive advantage

There is evidence to support that antler size influences mate selection in the red deer, and has a heritable component. Despite this, a 30-year study showed no shift in the median size of antlers in a population of red deer. [14] The lack of response could be explained by environmental covariance, meaning that lifetime breeding success is determined by an unmeasured trait which is phenotypically correlated with antler size but for which there is no genetic correlation of antler growth. [14] Alternatively, the lack of response could be explained by the relationship between heterozygosity and antler size, which states that males heterozygous at multiple loci, including MHC loci, have larger antlers. [15] The evolutionary response of traits that depend on heterozygosity is slower than traits that are dependent on additive genetic components and thus the evolutionary change is slower than expected. [15] A third possibility is that the costs of having larger antlers (resource use, and mobility detriments, for instance) exert enough selective pressure to offset the benefit of attracting mates; thereby stabilizing antler size in the population.

Protection against predation

If antlers functioned only in male–male competition for mates, the best evolutionary strategy would be to shed them immediately after the rutting season, both to free the male from a heavy encumbrance and to give him more time to regrow a larger new pair. Yet antlers are commonly retained through the winter and into the spring, [16] suggesting that they have another use. Wolves in Yellowstone National Park are 3.6 times more likely to attack individual male elk without antlers, or groups of elk in which at least one male is without antlers. [16] Half of all male elk killed by wolves lack antlers, at times in which only one quarter of all males have shed antlers. These findings suggest that antlers have a secondary function in deterring predation.

Snow shoveling for feeding

Reindeer Kebnekaise valley, Sweden (2007) 20070818-0001-strolling reindeer.jpg
Reindeer Kebnekaise valley, Sweden (2007)

Reindeer (genus Rangifer, whose sole member species R. tarandus comprises several distinctive subspecies of reindeer and caribou) use their antlers to clear away snow so they can eat the vegetation underneath. This is one possible reason that females of this species evolved antlers. [7] Another possible reason is for female competition during winter foraging. [13] Male and female reindeer antlers differ in several respects. Males shed their antlers prior to winter, while female antlers are retained throughout winter. [17] Also, female antler size plateaus at the onset of puberty, around age three, while males' antler size increases during their lifetime. [18] This likely reflects the differing life history strategies of the two sexes, where females are resource limited in their reproduction and cannot afford costly antlers, while male reproductive success depends on the size of their antlers because they are under directional sexual selection. [18]

Antenna for hearing

A six-year old moose undergoing domestication at Kostroma Moose Farm Luchik-the-Moose-and-Dr-Minaev-hp3188.jpg
A six-year old moose undergoing domestication at Kostroma Moose Farm

In moose, antlers may act as large hearing aids. Equipped with large, highly adjustable external ears, moose have highly sensitive hearing. Moose with antlers have more sensitive hearing than moose without, and a study of trophy antlers with an artificial ear confirmed that the large flattened (palmate) antler behaves like a parabolic reflector. [20]

Diversification

The diversification of antlers, body size and tusks has been strongly influenced by changes in habitat and behavior (fighting and mating). [13]

Capreolinae

Cervinae

Homology and evolution of tines

Antlers originated once in the cervid lineage. [21] The earliest fossil remains of antlers that have been found are dated to the early Miocene, about 17 million years ago. These early antlers were small and had just two forks. [21] As antlers evolved, they lengthened and gained many branches, or tines, becoming more complex. [21] The homology of tines has been discussed since the 1900s and has provided great insight into the evolutionary history of the Cervidae family. [22] [23] [24]

Recently, a new method to describe the branching structure of antlers was developed. [25] It is by using antler grooves, which are formed on the surface of antlers by growth, projecting the branching structure on the burr circumference, and making diagrams. Comparing the positional order among species on the diagram, the tine on the same position is homologous. The study revealed that three-pointed structures of Capreolinae and Cervini are homoplasious, and their subclades gained synapomorphous tines.

Exploitation by other species

Ecological role

Discarded antlers represent a source of calcium, phosphorus and other minerals and are often gnawed upon by small animals, including squirrels, porcupines, rabbits and mice. This is more common among animals inhabiting regions where the soil is deficient in these minerals. Antlers shed in oak forest inhabited by squirrels are rapidly chewed to pieces by them. [26] [27]

Trophy hunting

Antlered heads are prized as trophies with larger sets being more highly prized. The first organization to keep records of sizes was Rowland Ward Ltd., a London taxidermy firm, in the early 20th century. For a time only total length or spread was recorded. In the middle of the century, the Boone and Crockett Club and the Safari Club International developed complex scoring systems based on various dimensions and the number of tines or points, and they keep extensive records of high-scoring antlers. [28] Deer bred for hunting on farms are selected based on the size of the antlers. [29]

Hunters have developed terms for antler parts: beam, palm, brow, bez or bay, trez or tray, royal, and surroyal. These are the main shaft, flattened center, first tine, second tine, third tine, fourth tine, and fifth or higher tines, respectively. [30] The second branch is also called an advancer.

In Yorkshire in the United Kingdom roe deer hunting is especially popular due to the large antlers produced there. This is due to the high levels of chalk in Yorkshire. The chalk is high in calcium which is ingested by the deer and helps growth in the antlers. [31]

Shed antler hunting

A shed moose antler in Denali National Park and Preserve, Alaska (2010) Moose Antlers in Denali National Park.JPG
A shed moose antler in Denali National Park and Preserve, Alaska (2010)

Gathering shed antlers or "sheds" attracts dedicated practitioners who refer to it colloquially as shed hunting, or bone picking. In the United States, the middle of December to the middle of February is considered shed hunting season, when deer, elk, and moose begin to shed. The North American Shed Hunting Club, founded in 1991, is an organization for those who take part in this activity. [26]

In the United States in 2017 sheds fetch around US$10 per pound, with larger specimens in good condition attracting higher prices. The most desirable antlers have been found soon after being shed. The value is reduced if they have been damaged by weathering or being gnawed by small animals. A matched pair from the same animal is a very desirable find but often antlers are shed separately and may be separated by several miles. Some enthusiasts for shed hunting use trained dogs to assist them. [32] Most hunters will follow 'game trails' (trails where deer frequently run) to find these sheds or they will build a shed trap to collect the loose antlers in the late winter/early spring.

In most US states, the possession of or trade in parts of game animals is subject to some degree of regulation, but the trade in antlers is widely permitted. [33] In the national parks of Canada, the removal of shed antlers is an offense punishable by a maximum fine of C$25,000, as the Canadian government considers antlers to belong to the people of Canada and part of the ecosystems in which they are discarded. [34]

Carving for decorative and tool uses

A German powder flask made from a red deer antler, c.1570. Wallace Collection, London (2010) Wallace CollectionDSCF7493.JPG
A German powder flask made from a red deer antler, c.1570. Wallace Collection, London (2010)

Antler has been used through history as a material to make tools, weapons, ornaments, and toys. [35] It was an especially important material in the European Late Paleolithic, used by the Magdalenian culture to make carvings and engraved designs on objects such as the so-called Bâton de commandements and the Bison Licking Insect Bite . In the Viking Age and medieval period, it formed an important raw material in the craft of comb-making. In later periods, antler—used as a cheap substitute for ivory—was a material especially associated with equipment for hunting, such as saddles and horse harness, guns and daggers, powder flasks, as well as buttons and the like. The decorative display of wall-mounted pairs of antlers has been popular since medieval times at least.[ citation needed ]

The Netsilik Inuit people made bows and arrows using antler, reinforced with strands of animal tendons braided to form a cable-backed bow. [36] Several Indigenous American tribes also used antler to make bows, gluing tendons to the bow instead of tying them as cables. An antler bow, made in the early 19th century, is on display at Brooklyn Museum. Its manufacture is attributed to the Yankton Sioux. [37]

Through history large deer antler from a suitable species (e.g. red deer) were often cut down to its shaft and its lowest tine and used as a one-pointed pickax. [38] [39]

Ceremonial roles

Antler headdresses were worn by shamans and other spiritual figures in various cultures, and for dances; 21 antler "frontlets" apparently for wearing on the head, and over 10,000 years old, have been excavated at the English Mesolithic site of Starr Carr. Antlers are still worn in traditional dances such as Yaqui deer dances and carried in the Abbots Bromley Horn Dance.[ citation needed ]

Dietary usage

In the velvet antler stage, antlers of elk and deer have been used in Asia as a dietary supplement or alternative medicinal substance for more than 2,000 years. [40] Recently, deer antler extract has become popular among Western athletes and body builders because the extract, with its trace amounts of IGF-1, is believed to help build and repair muscle tissue; however, one double-blind study did not find evidence of intended effects. [41] [42]

Elk, deer, and moose antlers have also become popular forms of dog chews that owners purchase for their pet canines.

Shed hunting with dogs

Dogs are sometimes used to find shed antlers. The North American Shed Hunting Dog Association (NASHDA) has resources for people who want to train their dogs to find shed antlers and hold shed dog hunting events.

Related Research Articles

Moose Genus of mammals belonging to the deer, muntjac, roe deer, reindeer, and moose family of ruminants

The moose or elk is a member of the New World deer subfamily and is the largest and heaviest extant species in the deer family. Most adult male moose have distinctive broad, palmate antlers; most other members of the deer family have antlers with a dendritic ("twig-like") configuration. Moose typically inhabit boreal forests and temperate broadleaf and mixed forests of the Northern Hemisphere in temperate to subarctic climates. Hunting and other human activities have caused a reduction in the size of the moose's range over time. It has been reintroduced to some of its former habitats. Currently, most moose occur in Canada, Alaska, New England, New York State, Fennoscandia, the Baltic states, Poland, Kazakhstan and Russia.

Ungulate Group of animals that use the tips of their toes or hooves to walk on

Ungulates are members of the diverse clade Ungulata which primarily consists of large mammals with hooves. These include odd-toed ungulates such as horses, rhinoceroses, and tapirs; and even-toed ungulates such as cattle, pigs, giraffes, camels, sheep, deer, and hippopotamuses. Cetaceans such as whales, dolphins, and porpoises are also classified as even-toed ungulates, although they do not have hooves. Most terrestrial ungulates use the hoofed tips of their toes to support their body weight while standing or moving.

Deer Family of mammals belonging to even-toed ungulates

Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer (caribou), white-tailed deer, the roe deer, and the moose. Male deer of all species as well as female reindeer, grow and shed new antlers each year. In this they differ from permanently horned antelope, which are part of a different family (Bovidae) within the same order of even-toed ungulates (Artiodactyla).

Even-toed ungulate Order of mammals

The even-toed ungulates are ungulates—hoofed animals—which bear weight equally on two of their five toes: the third and fourth. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, odd-toed ungulates bear weight on an odd number of the five toes. Another difference between the two is that even-toed ungulates digest plant cellulose in one or more stomach chambers rather than in their intestine as the odd-toed ungulates do.

Reindeer Species of deer, also known as caribou

The reindeer, also known as the caribou in North America, is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, boreal, and mountainous regions of northern Europe, Siberia, and North America. This includes both sedentary and migratory populations. It is the only representative of the genus Rangifer. Herd size varies greatly in different geographic regions.

Chronic wasting disease Prion disease affecting the deer family

Chronic wasting disease (CWD), sometimes called zombie deer disease, is a transmissible spongiform encephalopathy (TSE) affecting deer. TSEs are a family of diseases thought to be caused by misfolded proteins called prions and include similar diseases such as BSE in cattle, Creutzfeldt-Jakob disease (CJD) in humans and scrapie in sheep. In the US, CWD affects mule deer, white-tailed deer, red deer, sika deer, elk, caribou, and moose. Natural infection causing CWD affects members of the deer family. Experimental transmission of CWD to other species such as squirrel monkeys, and genetically modified mice has been shown.

Rut (mammalian reproduction) Mating season of ruminant mammals

The rut is the mating season of certain mammals, which includes ruminants such as deer, sheep, camels, goats, pronghorns, bison, giraffes and antelopes, and extends to others such as skunks and elephants. The rut is characterized in males by an increase in testosterone, exaggerated sexual dimorphisms and increased aggression and interest in females. The males of the species may mark themselves with mud, undergo physiological changes or perform characteristic displays in order to make themselves more visually appealing to the females. Males also use olfaction to entice females to mate using secretions from glands and soaking in their own urine.

Red deer Species of mammal

The red deer is one of the largest deer species. A male red deer is called a stag or hart, and a female is called a hind. The red deer inhabits most of Europe, the Caucasus Mountains region, Anatolia, Iran, and parts of western Asia. It also inhabits the Atlas Mountains in Morocco and Tunisia, being the only species of deer to inhabit Africa. Red deer have been introduced to other areas, including Australia, New Zealand, the United States, Canada, Peru, Uruguay, Chile and Argentina. In many parts of the world, the meat (venison) from red deer is used as a food source.

Irish elk Extinct species of deer

The Irish elk also called the giant deer or Irish deer, is an extinct species of deer in the genus Megaloceros and is one of the largest deer that ever lived. Its range extended across Eurasia during the Pleistocene, from Ireland to Lake Baikal in Siberia. The most recent remains of the species have been carbon dated to about 7,700 years ago in western Russia.

<i>Megaloceros</i> Extinct genus of mammals in the family Cervidae

Megaloceros is an extinct genus of deer whose members lived throughout Eurasia from the early Pleistocene to the beginning of the Holocene and were important herbivores during the Ice Ages. The largest species, Megaloceros giganteus, vernacularly known as the "Irish elk" or "Giant elk", is also the best known. Fallow deer are thought to be their closest living relatives. Megaloceros is part of the deer family which includes moose, elk, reindeer, and other cervids.

Horn (anatomy)

A horn is a permanent pointed projection on the head of various animals that consists of a covering of keratin and other proteins surrounding a core of live bone. Horns are distinct from antlers, which are not permanent. In mammals, true horns are found mainly among the ruminant artiodactyls, in the families Antilocapridae (pronghorn) and Bovidae. Cattle horns arise from subcutaneous connective tissue and later fuse to the underlying frontal bone.

Sambar deer Species of deer

The sambar is a large deer native to the Indian subcontinent, South China, and Southeast Asia that is listed as a vulnerable species on the IUCN Red List since 2008. Populations have declined substantially due to severe hunting, local insurgency, and industrial exploitation of habitat.

Chital Species of deer

The chital, also known as spotted deer, chital deer, and axis deer, is a species of deer that is native to the Indian subcontinent. It was first described by German naturalist Johann Christian Polycarp Erxleben in 1777. A moderate-sized deer, male chital reach nearly 90 cm (35 in) and females 70 cm (28 in) at the shoulder. While males weigh 30–75 kg (66–165 lb), the lighter females weigh 25–45 kg (55–99 lb). It is sexually dimorphic; males are larger than females, and antlers are present only on males. The upper parts are golden to rufous, completely covered in white spots. The abdomen, rump, throat, insides of legs, ears, and tail are all white. The antlers, three-pronged, are nearly 1 m (3.3 ft) long.

Velvet antler

Velvet antler is the whole cartilaginous antler in a precalcified growth stage of the Cervidae family including the species of deer such as elk, moose, and caribou. Velvet antler is covered in a hairy, velvet-like "skin" known as velvet and its tines are rounded, because the antler has not calcified or finished developing. Velvet antler preparations are sold in China as part of traditional Chinese medicine, and in the United States and some other countries as a dietary supplement.

Thorolds deer Species of mammal

Thorold's deer is a threatened species of deer found in grassland, shrubland, and forest at high altitudes in the eastern Tibetan Plateau. It is also known as the white-lipped deer for the white patches around its muzzle.

Pecora Infraorder of mammals

Pecora is an infraorder of even-toed hoofed mammals with ruminant digestion. Most members of Pecora have cranial appendages projecting from their frontal bones; only two extant genera lack them, Hydropotes and Moschus. The name “Pecora” comes from the Latin word pecus, which means “horned livestock”. Although most pecorans have cranial appendages, only some of these are properly called “horns”, and many scientists agree that these appendages did not arise from a common ancestor, but instead evolved independently on at least two occasions. Likewise, while Pecora as a group is supported by both molecular and morphological studies, morphological support for interrelationships between pecoran families is disputed.

Elk Large antlered species of deer from North America and East Asia

The elk, also known as the wapiti, is one of the largest species within the deer family, Cervidae, and one of the largest terrestrial mammals in North America, as well as Central and East Asia. It is often confused with the larger Alces alces, which is called moose in North America, but called elk in British English, and related names in other European languages. The name "wapiti" is used in Europe for Cervus canadensis. It originates from the Shawnee and Cree word waapiti, meaning 'white rump'.

Deer hunting

Deer hunting is hunting for deer for meat or sport, an activity which dates back tens of thousands of years. Venison, the name for deer meat, is a nutritious and natural food source of animal protein that can be obtained through deer hunting. There are many different types of deer around the world that are hunted for their meat.

Capreolinae Subfamily of mammals

The Capreolinae, Odocoileinae, or the New World deer are a subfamily of deer. Alternatively, they are known as the telemetacarpal deer, due to their bone structure being different from the plesiometacarpal deer subfamily Cervinae. The telemetacarpal deer maintain their distal lateral metacarpals, while the plesiometacarpal deer maintain only their proximal lateral metacarpals. The Capreolinae are believed to have originated in the Middle Miocene, between 7.7 and 11.5 million years ago, in central Asia.

<i>Rucervus</i> Genus of mammals belonging to the deer, muntjac, roe deer, reindeer, and moose family of ruminants

Rucervus is a genus of deer from India, Nepal, Indochina, and the Chinese island of Hainan. The only extant representatives, the barasingha and Eld's deer, are threatened by habitat loss and hunting, and another species became extinct in 1938. The species of the genus Rucervus are characterized by a specific antler structure: its basal ramification is often supplemented with an additional small prong, the middle tine is never present, while the crown tines are inserted on the posterior side of the beam and may be bifurcated or fused into a small palmation.

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