Antler

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Mature red deer stag, Denmark Red deer stag 2009 denmark.jpg
Mature red deer stag, Denmark
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. [2]

Contents

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 muntjacs have small antlers and tusks. The musk deer, which are not true cervids, also bear tusks in place of antlers. [7]

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 [8] (the same material as human fingernails and toenails).

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. [9] The "horns" of a pronghorn (which is not a cervid but a antilocaprid) meet some of the criteria of antlers, but are not considered true antlers because they contain keratin. [10]

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, [11] and grow faster than any other mammal bone. [12] 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. [13]

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. [14] 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.[ citation needed ]

A 2019 study published in Science identified eight genes active in antler formation that are normally associated with bone cancer, particularly osteosarcoma. Additional tumor-suppressing and tumor-growth-inhibiting genes were determined to be responsible for regulating antler growth. This was taken to indicate that antler formation is more similar to a highly controlled form of cancer growth than to normal bone development. [15]

Antlers function as both weapons in male-male competition and as displays of sexual ornaments for females. [12] [16] Because mature antlers are no longer living during combat, antler fractures are incapable of being repaired following competition. A study in 2019 hypothesized that the periodic casting and regrowth of antlers might have evolved as a way to ensure the availability of complete antler sets to display each year. [17] Antler regeneration in male deer ensures that every mating season begins on a clean slate, as an increase in branching size and complexity happens each regeneration cycle in an individual. [14]

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. [11] 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. [11] 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. [11] 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. [18]

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. [19] 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. [19] 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. [20] 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. [20] 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, [21] 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. [21] 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.

Female antlers in reindeer

Reindeer in Kebnekaise valley, Sweden 20070818-0001-strolling reindeer.jpg
Reindeer in Kebnekaise valley, Sweden

Reindeer (Rangifer tarandus) are the only cervid species that inhabit the Arctic and subarctic regions of the globe, yet their most striking distinction is the presence of pedicles after birth and antlers in both males and females. [22] [23] One possible reason that females of this species evolved antlers is to clear away snow so they can eat the vegetation underneath. [7] Another possible reason is for female competition during winter foraging. [18] Espmark (1964) observed that the presence of antlers on females is related to the hierarchy rank and is a result of the harsh winter conditions and the female dominated parental investment. [24] Males shed their antlers prior to winter, while female antlers are retained throughout winter. [25] Also, female antler size plateaus at the onset of puberty, around age three, while males' antler size increases during their lifetime. [26] 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. [26] In other species of deer, the presence of antlers in females indicates some degree of intersex condition, the frequency of which has been seen to vary from 1.5% [27] to 0.02% [28] .

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. [30]

Diversification

The diversification of antlers, body size and tusks has been strongly influenced by changes in habitat and behavior (fighting and mating). [18]

Capreolinae

Cervinae

Homology and evolution of tines

Antler phylogenetics Antler phylogenetics - Samejima et al 2020.png
Antler phylogenetics

Antlers originated once in the cervid lineage. [31] 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. [31] As antlers evolved, they lengthened and gained many branches, or tines, becoming more complex. [31] The homology of tines has been discussed since the 1900s and has provided great insight into the evolutionary history of the Cervidae family. [32] [33] [34]

Recently, a new method to describe the branching structure of antlers was developed. [35] 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. [36] [37]

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. [38] Deer bred for hunting on farms are selected based on the size of the antlers. [39]

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. [40] 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. [41]

Shed antler hunting

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. [36]

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. [42] 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. [43] 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. [44]

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. [45] 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, an Inuit group, made bows and arrows using antler, reinforced with strands of animal tendons braided to form a cable-backed bow. [46] 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. [47]

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. [48] [49]

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. [50] 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. [51] [52]

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) [53] has resources for people who want to train their dogs to find shed antlers and hold shed dog hunting events.

Related Research Articles

<span class="mw-page-title-main">Moose</span> Mammal belonging to the deer family of ruminants

The moose or elk is the only species in the genus Alces. The moose is the tallest, and the second-largest, land animal in North America, only falling short of the American bison, in terms of mass. It is the world's tallest, largest and heaviest extant species of deer. Most adult male moose have broad, palmate antlers; other members of the deer family have antlers with a dendritic ("twig-like"), pointed configuration. Moose inhabit the circumpolar boreal forests or temperate broadleaf and mixed forests of the Northern Hemisphere, thriving in cooler, temperate areas as well as subarctic climates.

<span class="mw-page-title-main">Deer</span> Family of mammals

A deer or true deer is a hoofed ruminant mammal of the family Cervidae. The two main groups of deer are the Cervinae, including muntjac, elk (wapiti), red deer, and fallow deer; and the Capreolinae, including reindeer (caribou), white-tailed deer, roe deer, and 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).

<span class="mw-page-title-main">Artiodactyl</span> Order of mammals

Artiodactyls are mammals belonging to the order Artiodactyla. Typically, they are ungulates which bear weight equally on two of their five toes: the third and fourth, often in the form of a hoof. The other three toes are either present, absent, vestigial, or pointing posteriorly. By contrast, most perissodactyls bear weight on an odd number of the five toes. Another difference between the two is that many artiodactyls digest plant cellulose in one or more stomach chambers rather than in their intestine as perissodactyls do. The advent of molecular biology, along with new fossil discoveries, found that cetaceans fall within this taxonomic branch, being most closely related to hippopotamuses. Some modern taxonomists thus apply the name Cetartiodactyla to this group, while others opt to include cetaceans within the existing name of Artiodactyla. Some researchers use "even-toed ungulates" to exclude cetaceans and only include terrestrial artiodactyls, making the term paraphyletic in nature.

<span class="mw-page-title-main">Reindeer</span> Species of deer

The reindeer or caribou is a species of deer with circumpolar distribution, native to Arctic, subarctic, tundra, boreal, and mountainous regions of Northern Europe, Siberia, and North America. It is the only representative of the genus Rangifer. More recent studies suggest the splitting of reindeer and caribou into six distinct species over their range.

<span class="mw-page-title-main">Chronic wasting disease</span> 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. Natural infection causing CWD affects members of the deer family. In the United States, CWD affects mule deer, white-tailed deer, red deer, sika deer, elk, caribou, and moose. Experimental transmission of CWD to other species such as squirrel monkeys and genetically modified mice has been shown.

<span class="mw-page-title-main">Musk deer</span> Genus of mammals

Musk deer can refer to any one, or all seven, of the species that make up Moschus, the only extant genus of the family Moschidae. Despite being commonly called deer, they are not true deer belonging to the family Cervidae, but rather their family is closely related to Bovidae, the group that contains antelopes, bovines, sheep, and goats. The musk deer family differs from cervids, or true deer, by lacking antlers and preorbital glands also, possessing only a single pair of teats, a gallbladder, a caudal gland, a pair of canine tusks and—of particular economic importance to humans—a musk gland.

<span class="mw-page-title-main">Red deer</span> Species of hoofed 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 of Northern Africa; being the only living 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.

<span class="mw-page-title-main">Irish elk</span> 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 radiocarbon dated to about 7,700 years ago in western Russia. Its antlers, which can span 3.5 metres (11 ft) across are the largest known of any deer. It is not closely related to either living species called the elk, with it being widely agreed that its closest living relatives are fallow deer (Dama).

<span class="mw-page-title-main">Sambar deer</span> 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.

<span class="mw-page-title-main">Chital</span> Species of deer

The chital or cheetal, also known as the spotted deer, chital deer and axis deer, is a deer species native to the Indian subcontinent. It was first described and given a binomial name by German naturalist Johann Christian Polycarp Erxleben in 1777. A moderate-sized deer, male chital reach 90 cm (35 in) and females 70 cm (28 in) at the shoulder. While males weigh 70–90 kg (150–200 lb), females weigh around 40–60 kg (88–132 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 long.

<span class="mw-page-title-main">Thorold's deer</span> 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.

<span class="mw-page-title-main">Barren-ground caribou</span> Subspecies of deer

The barren-ground caribou is a subspecies of the reindeer that is found in the Canadian territories of Nunavut and the Northwest Territories, in northern Alaska and in south-western, Greenland. It includes the Porcupine caribou of Yukon and Alaska. The barren-ground caribou is a medium-sized caribou, smaller and lighter-colored than the boreal woodland caribou, with the females weighing around 90 kg (200 lb) and the males around 150 kg (330 lb). However, on some of the smaller islands, the average weight may be less. The large migratory herds of barren-ground caribou take their names from the traditional calving grounds, such as the Ahiak herd, the Baffin Island herds, the Bathurst herd, the Beverly herd, the Bluenose East herd, the Bluenose West herd, the Porcupine herd and the Qamanirjuaq herd.

<i>Cervus</i> Genus of deer and elk

Cervus is a genus of deer that primarily are native to Eurasia, although one species occurs in northern Africa and another in North America. In addition to the species presently placed in this genus, it has included a whole range of other species now commonly placed in other genera. Additionally, the species-level taxonomy is in a state of flux.

<span class="mw-page-title-main">Elk</span> Large antlered species of deer from North America and East Asia

The elk, or wapiti, is the second largest species within the deer family, Cervidae, and one of the largest terrestrial mammals in its native range of North America and Central and East Asia. The word "elk" originally referred to the European variety of the moose, Alces alces, but was transferred to Cervus canadensis by North American colonists. The name "wapiti" derives from a Shawnee and Cree word meaning "white rump" for the distinctive light fur in the rear region, just like the Bighorn Sheep.

<span class="mw-page-title-main">Reindeer hunting in Greenland</span> The practice of hunting reindeer for their meat, fur, and antlers in Greenland

Reindeer hunting in Greenland is of great importance to the Greenlandic Inuit and sports hunters, both residents and tourists. Reindeer (caribou) are an important source of meat, and harvesting them has always played an important role in the history, culture, and traditions of the Greenlandic Inuit. Controlled hunting is important for the welfare of reindeer, the quality of life for Inuit, both as food, and part of their culture and Greenlandic culture in general, and the preservation of tundra grazing areas. Therefore, scientific research is regularly performed to determine the quotas needed to maintain a proper ecological balance.

<span class="mw-page-title-main">Capreolinae</span> 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.

<span class="mw-page-title-main">American mountain deer</span> Extinct species of deer

Odocoileus lucasi, known commonly as the American mountain deer, is an extinct species of North American deer.

<span class="mw-page-title-main">Finnish forest reindeer</span> Subspecies of deer

The Finnish forest reindeer(Rangifer fennicus fennicus, also known as Eurasian or European forest reindeer is a rare subspecies of the reindeer native to Finland and northwestern Russia. They are found primarily in Russian Karelia and the provinces of North Karelia, Savonia and Kainuu in Finland, though some range into central south Finland. They are distinct from the semi-domesticated mountain reindeer in their larger size, longer legs and preference for dense boreal forest habitat, where they are rarely seen by humans, over the open tundra. The Finnish herd migrates seasonally back and forth across the long Russo-Finnish border.

<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 or swamp deer and Eld's deer, are threatened by habitat loss and hunting; another species, Schomburgk’s deer, went extinct in 1938. Deer species found within the genus Rucervus are characterized by a specific antler structure, where the basal ramification is often supplemented with an additional small prong, and the middle tine is never present. The crown tines are inserted on the posterior side of the beam and may be bifurcated or fused into a small palmation.

<span class="mw-page-title-main">Svalbard reindeer</span> Species of deer

The Svalbard reindeer is a small subspecies or species of reindeer found on the Svalbard archipelago of Norway. Males average 65–90 kg (143–198 lb) in weight, females 53–70 kg (117–154 lb), while for other reindeer generally body mass is 159–182 kg (351–401 lb) for males and 80–120 kg (180–260 lb) for females.

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PD-icon.svg This article incorporates text from a publication now in the public domain :  Chambers, Ephraim, ed. (1728). "antler". Cyclopædia, or an Universal Dictionary of Arts and Sciences . Vol. 1 (1st ed.). James and John Knapton, et al. p. 113.

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