Rut (mammalian reproduction)

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Male impalas fighting during the rut or breeding season Fighting impalas edit2.jpg
Male impalas fighting during the rut or breeding season

The rut (from the Latin rugire, meaning "to roar") 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, increased aggression, and increased interest in females. [1] 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. [1] [2] Males also use olfaction to entice females to mate using secretions from glands and soaking in their own urine. [1] [3] [4] Deer will also leave their own personal scent marking around by urinating down their own legs with the urine soaking the hair that covers their tarsal glands. Male deer do these most often during breeding season.[ citation needed ]

Contents

During the rut (known as the rutting period and in domestic sheep management as tupping ), males often rub their antlers or horns on trees or shrubs, fight with each other, wallow in mud or dust, self-anoint, and herd estrus females together. These displays make the male conspicuous and aid in mate selection.

The rut in many species is triggered by shorter day lengths. For different species, the timing of the rut depends on the length of the gestation period (pregnancy), usually occurring so the young are born in the spring. This is shortly after new green growth has appeared thereby providing food for the females, allowing them to provide milk for the young, and when the temperatures are warm enough to reduce the risk of young becoming hypothermic.

Cervidae

White-tailed deer

The rut for white-tailed deer (Odocoileus virginianus) usually lasts three weeks in the Northern Hemisphere and may occur most of the year in tropical zones. The rut is the time when white-tail deer, especially bucks, are more active and less cautious than usual. This makes them easier to hunt, as well as more susceptible to being hit by motor vehicles. [5]

Outdoors writer Charles Alsheimer has done research demonstrating the white-tailed deer rut is also controlled by the lunar phase and that the rut peaks seven days after the second full moon during October and November (the rutting moon), [6] while elk begin rutting during the September equinox on 21 September.

A white-tail doe may be in estrus for up to 72 hours and may come into estrus up to seven times if she does not mate. Cows may come into estrus up to four or more times if they do not mate. Some people believe that it is possible that does can enter a second estrus around 28 to 30 days from its first estrus if the doe isn't bred. There are also studies that show that some does actively participate in seeking out possible mates in areas where there are less males to increase the chances of finding males and being bred. [7]

The rut can start as early as the end of September and can last all the way through the winter months. Bucks usually begin this process when the velvet is falling off their antlers, and it can last all the way until they start to shed their antlers. The peak of the rut, however, is right in the middle. The average peak day for the white-tail rut in the U.S. is November 13. [8] Around this period of time, the bucks and does are very active, with the rut in full swing. For a hunter sitting in a tree stand at this time of the year, it is not uncommon to see many deer pass through his specific area, due to other deer chasing others.

White-tailed bucks in late rut in the Great Smoky Mountains Cades-cove-deer-fighting-tn1.jpg
White-tailed bucks in late rut in the Great Smoky Mountains

There are many behaviors a buck will exhibit during the rut. During pre-rut, bucks will spar with each other. Sparring is low-intensity aggressive behavior, involving mostly pushing and shoving. Bucks of different sizes will do this to each other. After pre-rut is finished, a buck will rub his antlers on a tree (thus making a "rub"), and make scrapes on the ground with his hooves: both of these are ways a buck will mark its territory and proclaim his dominance for other bucks to see. These activities are usually done at night. [9] Bucks will make many different scent markings using different scent glands. These scent glands include pedal glands found between the toes, preorbital glands found in the corners of their eyes, tarsal glands found on the lower inside of their hind legs, and the metatarsal glands found in the outside hind leg between ankle and hoof. The deer spread scent from these glands by rubbing hooves during ground scrapes, rubbing faces on tree limbs, and urinating down legs. [10]

The most prominent behavior of all during the heat of the rut is fighting, where bucks show their true dominance to others. In fighting, bucks usually battle against similar-sized deer, and small bucks do not normally challenge mature large ones: more often than not, smaller bucks fear the more mature bucks and leave or avoid the dominant deer's territory. These fights can be long in duration with the winner getting the group of does. Some fights result in injuries with some resulting in death.

Studies show that males of all age-classes increased their search efforts for mates during rut. According to the source, "One-year-old males seemed to invest less effort searching for females because movement rates (m/h) were 25–30% lower than in older males". This data indicates that the youngest males don't try as hard as older males to find a mate. [11] The energy expenditure of chasing and fighting during the breeding season can result in a buck losing an immense amount of weight, with some research documenting losses of as much as 20% of body weight with some sources showing body mass losses of up to 30%. [11] On average, a buck before breeding season can weigh up to 180 pounds (82 kg). After he has gone through the stages of the rut, he can lose about 50 pounds (23 kg) of weight, which is quite large, especially for only a few months of time. In the post-rut, a buck will need to replenish his body and catch up on the weight and energy he has lost.

Sources [12] [ non-primary source needed ] have stated that after the rut, a buck will go to a bedding spot and will remain "motionless" for a large amount of time, even to the extent of about two days, as he is thoroughly exhausted. After he has rested, he will get up and start to feed extensively, trying to catch up on all the nutrients his body requires. Croplands have much high carbohydrate grain in them, and a buck can be found here often, eating and getting nutrients. When the climate is extremely cold, a buck will sometimes resort to swamps and bogs, because of the warmer temperatures these areas hold.

Elk (Wapiti)

The timing of the elk rut depends on where they live. In the Northern Hemisphere, it occurs between mid-August and mid-October. In the Southern Hemisphere it occurs between mid-February and mid-April.

The rut tends to last somewhere between 20 and 45 days. [13] This varies on latitude, which affects the timing of spring and autumn and which can give elk a longer calving season and a longer rut. [14] During the rut, elk frequently use areas around fresh water, and tend to bed in heavy timber five to six hours per day. [15] :579 A cow elk will remain in estrus for 12 to 15 hours, if they are not bred during this time frame they will normally have another estrus cycle 18 to 28 days later. [14]

Elk use several different vocalizations during the rut. Some are made only by a certain sex or age class, and each is used for a different reason. The first of which being the cohesion call which is made by both sexes of elk, and is used to locate one another. [16] :225 An alarm squeal is made by both sexes of elk when they are on alert, during the rut these are used frequently by young bulls (male elks) being run off by the herd bull. [16] :228 Satellite bulls frequently spar with one another during the rut, and in turn make sparring squeaks. [16] :228

Bugling

A bugle is a vocalization made exclusively by bulls. The typical bugle consists of three acoustic parts, a low frequency "on-glide" that sounds guttural in tone, which then ascends into the highest frequency part of the call termed the "whistle", and the last portion of the call, the "off-glide" that returns to a low-frequency tone. [17] The function behind this acoustic structure of the bugle is directly correlated to the male's physiology and how different frequencies travel through varied environments. [18] [19] In terms of physiology, the larger an animal gets the lower frequency of sound it is able to produce. [20] This is because with an increase in size comes an increase in vocal fold length, and longer vocal folds confer an increased ability to produce lower frequency sounds. [20] Because of this relationship, a bugle can be directed toward other bulls or toward cows to demonstrate the size and thus fitness of the bull vocalizing. A bull will direct his bugle toward his cows while gathering them or while chasing an estrus cow. A herd bull will direct his bugle toward another bull to express his dominance over the herd, while a satellite bull may use his bugle to challenge the herd bull. [16] :229 The reason for the high-frequency portion of the bugle is due to the propagative efficiency of differing frequencies through varying environments. Studies have shown that as a bull's harem increases in diameter, meaning the cows become more dispersed, he tends to vocalize more frequently than if they were within closer proximity. [21] The higher-pitched section of the call propagates through the environment better, which is why the bull uses it to congregate a harem that is becoming more spatially dispersed and thus harder to defend. [21] Acoustic analyses comparing bull elk bugles with cow elk cohesion calls show a notable degree of acoustic similarity, indicating that both vocalizations may perform a congregating function, which is why the bugle is often used by the bull to condense his harem. [21]

Bugling physiology

The ability to produce such a high-frequency vocalization by such a large animal is unusual. [22] As explained above, this is because larger body mass positively correlates with longer vocal folds and thus lower frequency emission. Larger body size also corresponds with a decreased ability to emit high frequency vocalizations. [22] Bull elk overcome this by a unique anatomical mechanism that produces sound using a different pathway than the vibrations of the vocal folds. [23] Bull elk constrict their supra laryngeal vocal tract, specifically in the nasal cavity in order to create a smaller opening for exhaled air to pass through. [23] As air moves through this opening it causes the tissues to vibrate and produce the high frequency sound waves which comprise the "whistle" portion of the bugle. [24] This anatomical development for bioacoustics in elk was discovered upon sonographic analysis of bugle vocalizations which revealed a biphonetic (two simultaneous frequencies) display. [23] One frequency was high (the result of the supra laryngeal constriction), and the other was low (the result of normal vocal fold oscillations). [23]

Yelping

Yelping also known as "grunting" is usually only made by herd bulls when they are excited. They are made more often while interacting with cows than with other bulls. "Yelping commonly was accompanied by contractions of the penile region with simultaneous emission of short spurts of urine." [16] :230

Phases

The rut has six phases: the pre-rut, the first breeding phase, the first rest phase, the second breeding phase, the second rest phase, and the third breeding phase. [14]

  • The pre-rut takes place from mid-August through the beginning of September. During the pre-rut bulls begin bugling and gathering their herds. Bulls will bugle to attract cows as well as to express dominance over other bulls. A "herd" bull is the dominant bull in a herd. Younger, smaller bulls are known as satellite bulls, as they tend to cling to the edges of a herd trying to pick up any cows willing to leave the herd. Larger satellite bulls will challenge the herd bull to try and take control of the herd. These challenges include a good deal of bugling as well as fighting. [14]
  • The first breeding phase of the rut takes place between the beginning and the middle of September. This is when the three year and older cows come into estrus. During this time herd bulls bugle to keep their cows close by, they also answer the bugles of satellite bulls to let them know they are still dominant. A herd bull will also bugle while approaching a cow in estrus so the cows become familiar with his bugles. [14]
  • The first rest phase of the rut occurs between the middle and the end of September. At this time the older cows are predominantly out of estrus and the younger cows have not yet come into estrus. During the rest period, satellite bulls will try to join the herd while the herd bull is resting.
  • The second breeding phase of the rut takes place three to four weeks after the first breeding phase. This is due to younger cows coming into estrus, as well as older cows that were not bred on their first estrus cycle coming back into estrus. Herd bulls are less aggressive towards satellite bulls at this phase in the rut due to exhaustion. [14] The second phase of the rut may have the most bugling activity due to the combination of the testosterone levels of the younger bulls rising, and the herd bull still trying to maintain control of the herd.
  • The second rest phase of the rut occurs around the middle of October. By this time the original herd bull usually does not have control of the herd, due to a great decline in physical condition. Terry Bowyer states, "Elk were observed feeding in the following percentages of observations: master bulls 24%; bachelor bulls 53%; yearling males 62%; cows 64%; and calves 62%" (Bowyer uses the terms "master bulls" and "bachelor bulls" which have the same meaning as "herd bulls" and "satellite bulls"). [15] :577 Herd bulls do not have time to feed during the rut due to constantly fighting other bulls as well as chasing and breeding cows. [14]
  • Occasionally a third breeding phase will occur. This will usually take place around the end of October or early November. This is a result of yearling cows coming into estrus for the first time or two-year-old cows coming into a second estrus cycle. Since most of the herd bulls have left the herd by this time of year, the breeding is usually done by the younger satellite bulls. After this phase the rut is over, most bulls will leave the cows and form bachelor herds to spend the winter with; however young bulls will usually remain with the cows throughout the winter. [14]

Other deer species

Red deer during the rut

Fallow deer

The fallow deer (Dama dama) is an ungulate which employs an unusual strategy for mating during the rut. This strategy is the creation of a lek, a display area presented to the females where the males gather and allow the females to choose a mate based upon their traits alone while reducing predation risk, disturbance to copulation, parasite transmission and the cost of looking for a mate. [25] [4] When females come to the lek they leave soon after mating but the males will tend to stay in the lek to court other females until the end of the rutting season. [26] However, male fallow deer which are unsuccessful in mating will leave the lek sooner than other males and they will adopt other strategies to compensate for their lack of mating success in the lek. Furthermore, the duration spent in the lek is positively correlated with the behavioral traits of male display frequency and aggression, male hierarchical position and secondary sex characteristics such as antler size. [27] Overall, lekking species such as the fallow deer have a short intense rutting season where the males face intrasexual competition, territory defense and management of females within their territory. [28]

Elephants

In elephants, the breeding season is less pronounced than in ungulates and it usually spikes when the rains season occurs or shortly thereafter. [29] [1] The rut is observed in both African and Asian elephants and it is referred to as musth. [30] Its meaning is derived from the Urdu word mast meaning intoxication. The most prominent characteristics of an elephant in rut are heightened sexual and aggressive activity along with copious temporal gland secretion and continual urine discharge. [1] [31] Also it has been observed that males will have a higher concentration of testosterone and an increased likelihood of associating with female groups during musth. Similarly to deer or mountain goats, elephants will tusk the ground throwing vegetation, logs and objects into the air and occasionally at subordinates. [1]

Moose

Moose have a series of rutting events that are similar to those seen in other deer species, however, they have several characteristic behaviors which give them a distinct rut. The first of these behaviors is a challenger gait where the bull moose will sway back and forth and circle the rival bull while dipping his antlers down. [32] Another typical behaviour seen in moose especially during the pre-rut stage is mock battling. This is a display meant to scare away other rival males where the bull moose will destroy trees and vegetation prior to engaging in a fight. [33] Also, a behaviour known as displacement feeding is observed in male moose and it refers to the hasty movements made by the moose while it is feeding as it keeps an intense gaze upon rival bull moose. [34] Furthermore, as seen in other deer species male moose will dig mud pits and soak them in urine and the females will fight over possession of these wallows. [34] The pre-rutting season typically begins during August and is marked by bull moose leaving the younger satellite bulls. [33] During this stage there is much mock fighting and the pre-rut ends in September when the bull moose emerge from the solidarity of heavily wooded areas. Then begins the searching stage of the rut where the male seeks the moose cow in estrus and the instances of displacement feeding and tension between rival males increases. [33] Once a potential mate has been found the male enters the display stage of the stage which lasts one to three days. During this time he will court the female by standing sidewise three to five yards from the female moose to show himself as a mate. If successful he will get to mate with her for several days and then move on to a new partner. This pattern of behaviours will then repeat with successive mates until late October or early November. Following the mating season, bull moose spend long hours resting and feeding before forming their usual winter groupings. [33]

Although the battle between males is the main contest, there is also a battle between females. Usually, this occurs between an older cow moose and a younger female. [35] The mature cow will attempt to stop the younger one from coming near to the wallow in a vicious attack with her forelegs and if the younger female gets to lie in the wallow the older female will drive her out of it only to return to lie in it and take up as much space as possible. [33] During this event the bull moose will not interfere and he will just watch in plain sight.

Other mammals

Further reading

See also

Notes

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 distinctively broad, palmate antlers; other members of the deer family have antlers with a dendritic ("twig-like"), pointed configuration. Moose are winter specialists, and typically 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">Lek mating</span> Type of animal mating behaviour

A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an available plot of space able to be utilized by displaying males to defend their own share of territory for the breeding season. A lekking species is characterised by male displays, strong female mate choice, and the conferring of indirect benefits to males and reduced costs to females. Although most prevalent among birds such as black grouse, lekking is also found in a wide range of vertebrates including some bony fish, amphibians, reptiles, and mammals, and arthropods including crustaceans and insects.

<span class="mw-page-title-main">Père David's deer</span> Species of mammals native to China

The Père David's deer, also known as the milu or elaphure, is a species of deer native to the subtropical river valleys of China. It grazes mainly on grass and aquatic plants. It is the only extant member of the genus Elaphurus. Some experts suggest demoting Elaphurus to a subgenus of Cervus. Based on genetic comparisons, Père David's deer is closely related to Eld's deer.

<span class="mw-page-title-main">European fallow deer</span> Species of hooved mammal

The European fallow deer, also known as the common fallow deer or simply fallow deer, is a species of ruminant mammal belonging to the family Cervidae. It is historically native to Turkey and possibly the Italian Peninsula, Balkan Peninsula, and the island of Rhodes in Europe. Prehistorically native to and introduced into a larger portion of Europe, it has also been introduced to other regions in the world.

<span class="mw-page-title-main">Muskox</span> Arctic land mammal

The muskox, also spelled musk ox and musk-ox, plural muskoxen or musk oxen, is a hoofed mammal of the family Bovidae. Native to the Arctic, it is noted for its thick coat and for the strong odor emitted by males during the seasonal rut, from which its name derives. This musky odor has the effect of attracting females during mating season. Its Inuktitut name "umingmak" translates to "the bearded one".

A bachelor herd is a herd of (usually) juvenile male animals who are still sexually immature or 'harem'-forming animals who have been thrown out of their parent groups but not yet formed a new family group. It may also refer to a group of males who are not currently territorial or mating with females.

<span class="mw-page-title-main">Musth</span> Condition in male elephants

Musth or must is a periodic condition in bull (male) elephants characterized by aggressive behavior and accompanied by a large rise in reproductive hormones.

<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">Common eland</span> Second largest antelope in the world

The common eland, also known as the southern eland or eland antelope, is a large-sized savannah and plains antelope found in East and Southern Africa. It is a species of the family Bovidae and genus Taurotragus. An adult male is around 1.6 m (5.2 ft) tall at the shoulder and can weigh up to 942 kg (2,077 lb) with a typical range of 500–600 kg (1,100–1,300 lb), 340–445 kg (750–981 lb) for females). It is the second-largest antelope in the world, being slightly smaller on average than the giant eland. It was scientifically described by Peter Simon Pallas in 1766.

<span class="mw-page-title-main">Common tsessebe</span> Subspecies of the subfamily Alcelaphinae in the family Bovidae

The common tsessebe or sassaby is the southern, nominate subspecies of Damaliscus lunatus, although some authorities have recognised it as an independent species. It is most closely related to the Bangweulu tsessebe, sometimes also seen as a separate species, less to the topi, korrigum, coastal topi and tiang subspecies of D. lunatus, and less to the bontebok in the same genus. Common tsessebe are found in Angola, Zambia, Namibia, Botswana, Zimbabwe, Eswatini, and South Africa.

<i>Phoca</i> Genus of carnivores

Phoca is a genus of the earless seals, within the family Phocidae. It now contains just two species, the common seal and the spotted seal. Several species formerly listed under this genus have been split into the genera Pusa, Pagophilus, and Histriophoca. Until recently, Phoca largha has been considered a subspecies of Phoca vitulina but now is considered its own species. For this reason, the fossil history of the genus is unclear, and it has formerly been used as wastebasket taxon for a number of fossils of uncertain affinity.

The estrous cycle is a set of recurring physiological changes induced by reproductive hormones in females of mammalian subclass Theria. Estrous cycles start after sexual maturity in females and are interrupted by anestrous phases, otherwise known as "rest" phases, or by pregnancies. Typically, estrous cycles repeat until death. These cycles are widely variable in duration and frequency depending on the species. Some animals may display bloody vaginal discharge, often mistaken for menstruation. Many mammals used in commercial agriculture, such as cattle and sheep, may have their estrous cycles artificially controlled with hormonal medications for optimum productivity. The male equivalent, seen primarily in ruminants, is called rut.

<i>Ovis</i> Genus of mammals

Ovis is a genus of mammals, part of the Caprinae subfamily of the ruminant family Bovidae. Its seven highly sociable species are known as sheep or ovines. Domestic sheep are members of the genus, and are thought to be descended from the wild mouflon of central and southwest Asia.

<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">California mule deer</span> Subspecies of deer

The California mule deer is a subspecies of mule deer whose range covers much of the state of California.

Elk farming is an agricultural industry for the production of elk as livestock or for the sport of hunting. Elk have a variety of uses. The velvet antler or, the antler in the premature stages of growth, is believed by some to have medicinal purposes. Elk are also raised for venison, their meat. All of these markets are rising in popularity thus causing an increase in the breeding industry. Other species of similar type are farmed in the same way such as deer, moose, and red stag.

Seasonal breeders are animal species that successfully mate only during certain times of the year. These times of year allow for the optimization of survival of young due to factors such as ambient temperature, food and water availability, and changes in the predation behaviors of other species. Related sexual interest and behaviors are expressed and accepted only during this period. Female seasonal breeders will have one or more estrus cycles only when she is "in season" or fertile and receptive to mating. At other times of the year, they will be anestrus, or have a dearth of their sexual cycle. Unlike reproductive cyclicity, seasonality is described in both males and females. Male seasonal breeders may exhibit changes in testosterone levels, testes weight, and fertility depending on the time of year.

<span class="mw-page-title-main">Self-anointing in animals</span> A behaviour whereby a non-human animal smears odoriferous substances over themselves

Self-anointing in animals, sometimes called anointing or anting, is a behaviour whereby a non-human animal smears odoriferous substances over themselves. These substances are often the secretions, parts, or entire bodies of other animals or plants. The animal may chew these substances and then spread the resulting saliva mixture over their body, or they may apply the source of the odour directly with an appendage, tool or by rubbing their body on the source.

<span class="mw-page-title-main">Hunting behavior of gray wolves</span>

Single wolves or mated pairs typically have higher success rates in hunting than do large packs; single wolves have occasionally been observed to kill large prey such as moose, bison and muskoxen unaided. This contrasts with the commonly held belief that larger packs benefit from cooperative hunting to bring down large game. The size of a wolf hunting pack is related to the number of pups that survived the previous winter, adult survival, and the rate of dispersing wolves leaving the pack. The optimal pack size for hunting elk is four wolves, and for bison a large pack size is more successful.

Bellow is a type of animal vocalization common amongst bulls and other large animals such as Rhinoceros, Bison, Buffalo, Yak And Red Deer. It's a form of roaring and reverberating sound. Also, the bull snake bellows and hisses as its defensive sounds, It bellows at first in a short period high amplitude, followed by a longer period of low amplitude before it maintains a constant sound.

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