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BEHAVIOUR - Editorial Comment

Editorial Comment (Editorial Overview Text Replicated on Overall Species page - Loxodonta africana - African Elephant)

Sexual behaviour in elephants appears to vary, with cows being described as actively soliciting males, but also as ignoring actual matings, continuing to feed. Females in oestrus may attract males by means of loud calls. Females may actively solicit a bull, e.g. backing into him, and there may be interactions such as short mock chases, head butting between the bull and cow, twining trunks, the male placing his trunk over the cow, etc. A female may choose one bull and remain close to that individual, and refuse to mate with other bulls. A male may check the oestrus status of the cow by touching her genital area with his trunk tip then placing the trunk tip in his mouth. Most guarding of oestrus cows by bulls is carried out by older bulls (35 years old or older). The dominant guarding bull may mate with the oestrus female perhaps three times over a 24 hour period. It has been variously suggested that matings by multiple bulls indicate an inexperienced cow who has not allied with a single bull, newly arrived dominant bulls taking over from bulls lower down in the hierarchy, or overcrowding of the population. It may also be that early in oestrus, when the cow has not yet ovulated (and is therefore not fertile) promiscuous matings may occur with minimal aggression, but late in oestrus the dominant bull prevents approach to the cow by other bulls. For the actual mating, the male approaches from behind and places his forelegs along the cow's back, lowering himself on his hind legs in a squatting position. The erect, S-shaped penis thrusts forward into the vulva, which is turned backwards by erection of the female's clitoris. The male then partially rises on his hind legs. There are no pelvis thrusts by the male but the penis moves inside the uro-genital canal. Actual copulation takes about one minute. 

Musth is not strictly speaking sexual behaviour; males do not need to be in musth in order to mate. However, musth may affect breeding success. A male in musth ranges over a larger area and is more likely to come into contact with females in oestrus. Males in musth gain dominance in the male hierarchy and are more likely to compete successfully for females in oestrus. Females may actively prefer a bull in musth to one not in musth. In addition to increased aggression, signs of musth include higher head carriage, high, widely spread carriage of the ears, urine dribbling, swelling and discharge from the temporal glands and uttering of the "musth rumble": low, pulsating growls; they also have greatly increased testosterone levels. Bulls in musth spend less time feeding than usual and lose body condition. Young bulls do not enter musth; in African elephants this may not be seen until bulls are at least 26 - 32 years old. Musth lasts longer in older than in younger bulls. There is some seasonality in when bulls come into musth, with peak occurrence in the rainy season. A given bull may come into musth at about the same time each year, although this may change as he gains or loses position in the hierarchy. It is possible that dominant bulls in musth may suppress the development of musth in lower ranking bulls.

(References are available in detailed literature reports below)

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Sexual Behaviour

Source Information
  • Elephants are polygynous. (B396.3.w3)
  • Females in oestrus give loud, low-frequency calls. These may attract males form several kilometers away. (B147)
    • These calls are largely inaudible to humans. (B147)
  • Females at Disney's Animal Kingdom, kept in an all female herd, showed the highest rate of low-frequency vocalisation and of repeated calling during the anovulatory follicular phase of the oestrus cycle, significantly higher than during the luteal phase. In the wild, this might attract males to the herd in this pre-ovulatory period. (J395.43.w1)
  • Females in oestrus tend to choose a large male in musth; they actively avoid most males. (B147)
  • It may be that an oestrus female chooses a male she as a long familiarity with. (B147)
  • The chosen male guards females from other males. (B147)
  • Females in oestrus emit a large infrasonic call which attracts bulls. (B285.w3)
  • Females in oestrus appear to prefer large bulls to small bulls and bulls in musth to bulls not in musth. (B285.w3)
  • During mid-oestrus, a female goes into consortship with a musth male; he guards her from the attentions of other lower-ranking bulls. Females prefer musth to non-musth males. (J334.37.w1)
  • An oestrus female chooses which bull to mate with and refuses to mate with a bull she does not want. (B285.w3)
  • Young, inexperienced, cows may not ally with a single bull, and may then be harassed and chased by multiple bulls during their period of heat. (B384.5.w5)
  • During the peak days of oestrus the female is guarded by a musth bull. (J396.22.w1)
  • Distinct behaviours may be seen in females approaching and during oestrus (described from females at Amboseli, Kenya): (B396.3.w3)
    • Wariness, with the female moving away from approaching bulls; (B396.3.w3)
    • Oestrous walk, in which the female, spotting an approaching male, walks briskly from her group, head high and held to one side, watching the bull following her, then walking in an arc and returning to her group. (B396.3.w3)
    • Chase, in which the female may run rather than walk, with one or more bulls pursuing her, usually in a wide arc back to the group - the female may stop if a bull touches her; (B396.3.w3)
    • Consortship, with the cow remaining in close proximity to a large bull for a short period, during which the female stays close to her bull and he threatens or chases off bulls which approach. (B396.3.w3)
  • Males approaching an oestrus cow check her reproductive condition by placing the trunk tip on her genital area and then into their mouths. (B384.5.w5)
  • Males show flehmen - testing the urine or urogenital orifice with the trunk tip then placing it inside the mouth to the vomeronasal organ - ten times as often with females in oestrus as with non-oestrus females. (B396.3.w3)
  • The dominant guarding bull generally mates with the oestrus female only about three times in 24 hours. (B384.5.w5)
  • Towards the end of the heat, the dominant bull loses interest. (B384.5.w5)
  • Nearly 90% of guarding behaviour by bulls is carried out by bulls of 35 years old or older. (B384.5.w5)
  • Females may bellow after being mated. (B384.5.w5)
  • The bull and cow may on occasion show behaviour such as head butting one another and twining trunks. Other behaviours which may occur include the male caressing the female with his trunk, placing the tip of his trunk in the cow's mouth and gently butting the cow. (B384.5.w5)
  • The female mates with the highest ranking bull at any time; this may involve several different bulls over the period of heat, since higher ranking bulls may arrive during this time. (B384.5.w5)
  • Observations of elephants in populations in East Africa found that, for several females in oestrus, while several bulls were attendant, the largest was clearly dominant to the others. (P17.21.w1)
  • Mating takes place in the usual position for quadrupeds, with the male mounting the female from behind. (B453.6.w6)
  • In undisturbed populations which are not overcrowded, it is probable that younger bulls give way to the largest bull, usually without any dispute or fighting. (B453.6.w6)
  • In overcrowded conditions, successive matings by different bulls have been observed. (B453.6.w6)
  • The dominant bull approaches the cow, caresses her with his trunk and the elephants may mock tussle, or place their trunks in one another's mouths. The bull then butts the cow on her hind quarters, in response to which she moves away, and this is repeated several times in a "mock chase", sometimes with inconclusive mounting attempts. Finally the cow stands still, the male places his forelegs on the cow's hind quarters, his buttocks being lowered almost to a squatting position. The penis is erect and curved concavely with the tip bent to give an S-shape as the bull thrusts forwards to the cow's vulva; the female's clitoris also erects, resulting in the vulva being moved down and backwards. The bull then partly rises on his hind legs with his forelegs extended along the cow's back and his tail held out behind him. (B453.6.w6)
  • Mating itself takes only about 45 seconds. (B384.5.w5)
  • Mating takes perhaps two minutes, with the bull placing his trunk along the female's back and standing up on his hind legs; the erect penis, S-shaped is placed into the opening of the urogenital canal, between the hind legs. (B396.3.w3)
  • Mounting, intromission and climax takes up to about 120 seconds. Copulation may be repeated at intervals of 10 to 20 minutes, with the bull and cow standing, resting or browsing in the intervening periods. (B453.6.w6)
  • A chase may or may not occur prior to mating. Immediately prior to mounting the cow, the bull lays his trunk and tusks along the cow's back. Copulation lasts less than a minute, with not pelvic thrusts from the male, but movement of the penis inside the uro-genital canal. (B451.4.w4)
    • Observation of captive elephants at Kronberg Zoo noted that the bull frequently touched his trunk to the female's vulva, as well as the occurrence of chases. (B451.4.w4)
  • In observations on one cow, in the early part of oestrus there were promiscuous matings and no aggression between bulls, however, later there was more aggression and fighting, ending with one large dominant bull remaining close to the cow and threatening other males if they approached. (B451.4.w4)
    • This may be related to the fact that ovulation occurs late in the oestrus period, so only matings during the late stage will be fertile. (B451.4.w4)
  • Initially, cows may retreat from courting bulls, or may engage in a pushing contest or head-on confrontation. A male courting a female lays his trunk over her back repeatedly. (B387.w4)
  • Several males may follow a cow in oestrus and there may be intense competition, even fighting. (B387.w4)
  • While there may be intense competition on some occasions, at other times mating occurs with minimal fuss and disruption. (B387.w4)
  • The female may solicit a bull, backing into him or rubbing her clitoris against him; there may then be a short chase, stopped when the bull touches the cow with his trunk. The male may also take the cow's tail in his mouth and butt the cow's hindquarters. (B387.w4)
  • For mating the cow places her hind legs in a wider stance and leans back. (B387.w4)
  • Several matings by one or different bulls may occur in succession. (B387.w4)
  • Copulation lasts 30 to 90 seconds. (B387.w4)
  • In some matings the cow continues feeding during the mating. (B387.w4)
  • A bull entering a family unit would usually sniff the genital area of each cow in turn to determine their reproductive condition. (B462.14.w14)
  • Bull elephants at Manyara were observed to run after an oestrus female, which would stop when he caught up and placed his trunk on her rump. The male then reared up onto his hind legs, straddling her back, or laid his head on her back and levered himself up, so the forefeet were either side of the female's spine. Mounting and pelvic thrusts lasted only about 30-40 seconds, sometimes with deep groans from the bull, but without obvious reaction from the female. The rest of the family unit showed great excitement during the chase and immediately after the mating, then all settled down. (B462.14.w14)
  • Matings are often followed by a "mating pandemonium" with the female's family rushing over, rumbling, screaming, trumpeting, urinating, defecating and producing temporal gland secretions. (J396.22.w1)

Musth:

  • Male elephants of about 25 years of age and older annually enter "musth". (B147)
  • Musth lasts only a few days or weeks in young elephants but in those 35 years old or older it lasts for two to five months. (B147)
  • Musth is not totally synchronised between males, but is particularly frequent during and immediately after the rainy season. (B147)
  •  A given male enters musth at about the same time each year. (B147)
  • Characteristically, elephants in musth: (B147)
    • Produce copious secretions from the temporal gland behind the eye;
    • Show continuous discharge of urine;
    • Show a great increase in aggressive behaviour;
    • Seek and associate with groups of female elephants. 

    (B147

  • Males in musth are dominant to other males and are usually able to defeat them, even if the other males are larger and are higher-ranking normally. (B147)
  • Bull elephants will breed with females whether or not they are in musth. (B10.49.w21)
  • Musth in African elephants is generally less pronounced than it is in Asian elephants. (B285.w3)
  • In Amboseli, Kenya, bulls only start to develop musth at about 29 years of age, when almost fully grown. (B285.w3)
  • Musth may last two to three months and generally occurs at the same time as periods of high rainfall. (B285.w3)
  • Musth bulls: carry their heads higher than normal, carry their ears high and wide spread, show swelling and discharge from the temporal glands and continuously dribble urine (containing pheromones). They also emit a characteristic "musth rumble" of low, pulsating growls. They are more likely to fight than are other bulls. They have a greatly reduced food intake and use fat reserves. (B285.w3)
  • Musth in the African elephant is characterised physically by continuous dribbling of urine, strong odour, a greenish colouration of the proximal penis and distal sheath, and pronounced enlargement of and copious secretion from the temporal glands. Behaviourally, musth bulls show increased aggression, are more sexually active, have a higher dominance rank than when not in must and are more often found in association with females. Musth males are found in consort relations with oestrus females significantly more often than are non-musth males. (J9.292.w1)
  • Musth bulls are able to dominate larger non-musth bulls who are larger and normally higher-ranking. Bulls in musth are highly aggressive, but escalated contests are rare. Signs of musth indicate the state of aggression. Males in musth spend less time eating and lose weight and this loss in condition is more severe when the period of musth is lengthened. Bulls come into musth asynchronously, with each male coming into musth at the same time each year, with gradual changes as the male rises or decreases in rank. (J334.37.w1)
  • Bulls have an increase in social status while in musth; a small bull in musth is likely to fight successfully against a larger bull which is not in musth. (B285.w3)
  • Musth bulls travel further than do bulls not in musth and may therefore be more likely to locate female elephants in oestrus. (B285.w3)
  • Musth occurs annually, peaks during the wet season, and may last from only a few days to several months. (B384.8.w8)
  • Signs of musth include swelling of and secretion from the temporal gland, urine dribbling and increased aggression. (B384.8.w8)
  • Bulls in musth spend less time than normal in feeding or resting, and lose body condition. They spend more time than usual in travelling and in interacting with conspecifics. (B384.8.w8)
  • Musth bulls may move long distances: 224 km in 22 days for one bull and 125 km in 30 days for another have been recorded. This increases the likelihood that they will encounter cows in heat. Additionally, since the bull may cover the ranges of several clans, outbreeding is more likely. (B384.8.w8)
  • Bulls in musth have increased testosterone levels: up to 50 times the level seen outside musth. (B384.8.w8)
    • In Addo Park, testosterone levels have been measured as 3.5 ng/mL for the lowest ranking non-musth bull to 19.8 ng/mL in a bull just coming out of musth. (B384.8.w8)
  • Musth bulls lose weight; this may be partially due to the reduced feeding and increased exercise, but may also be associated with the increased testosterone levels producing a higher metabolic rate. (B384.8.w8)
  • Bulls in musth hold the head high and the ears out and high; movement is also changed and displays of aggression such as kneeling and tusking the ground, or throwing objects, are seen. (B384.8.w8)
  • African musth bulls produce a low, pulsating call, at frequencies as low as 14 Hz and a volume up to 108 decibels, known as the musth rumble. This call is made most frequently when the bull is alone, less often when he is with cows. Ear flapping or waving occurs at the same time as the call and a loud ear flap occurs as the call ends. Older musth bulls call more often than do younger ones. Cows near the bull will reply with low frequency calls of their own. (B384.8.w8)
  • Musth may be seen year round, but there are recognised seasonal occurrences; in southern Africa it appears more seasonal than in eastern Africa. (B384.8.w8)
  • In a given bull, musth occurs at about yearly intervals, although it may not occur if the individual lost condition severely during the previous musth. (B384.8.w8)
    • The time at which a bull comes into musth may change as his position in the bull hierarchy improves, with musth then occurring closer to when the most cows are in heat, and as his position declines, when musth may become shorter and more sporadic. (B384.8.w8)
    • In African elephants, musth may first appear when a bull is 24 years old, but usually first occurs at 26 to 32 years. The oldest bulls come into musth first, before even encountering any cows in heat, while males of 35-40 years develop musth only after several weeks with cow groups and longer still is required for bulls under 35 to come into musth and then they only remain in musth for a few days. (B384.8.w8)
    • Bulls do not compete successfully for cows until they are about 30 years old. (B384.8.w8)
    • Musth bulls out-compete older, larger bulls which are not in musth. (B384.8.w8, B396.3.w3)
    • Dominant bulls in musth may suppress the development of musth in lower ranking bulls. (B384.8.w8)
  • In Amboseli National Park, Kenya, clear instances of musth have been described; these bulls dominate matings and also show aggression as well as dribbling from the penis, which becomes greenish. (B451.4.w4)
  • Musth involves intense sexual interest in females in oestrus, intense aggression towards other bulls, secretion of fluid from the temporal gland and may include constant dribbling of urine. Swelling of the temporal gland and secretion of musth fluid from the glands clearly indicates the onset of musth. The swelling becomes larger, the fluid more viscous, pungent and of greater volume as musth progresses; urine dribbling starts at this stage and the urine has a distinct odour; the sheath and penis become stained green; they also urinate with the penis inside the sheath, spraying urine onto their hind legs. Musth bulls rub their temporal glands on trees more often than when not in musth, have a characteristic walk with the head held high and ears spread, and may wave the ears one at a time, thrusting the ear forward suddenly (this has only been observed in African, not Asian, bulls in musth). A distinct musth rumble is also recognised; African savannah elephant bulls may call at up to 108 decibels (measured at 1 m distance) and with a frequency as low as 14 Hz. (B396.3.w3)
  • Musth gradually increases in frequency, intensity and duration as bulls get older. Not all males come into musth at the same time, but in areas where breeding is seasonal, the greatest proportions of bulls in musth are noted during the main mating season. It has also been noted that the largest most dominant bulls tend to be in musth during the best periods for mating. (B396.3.w3)
  • Males in musth utter significantly more calls when they are alone and apparently searching for females than when they are in association with females. The musth rumbles can carry quite long distances and may assist males to avoid unexpectedly meeting other high-ranking aggressive males while wandering in search of females in oestrus. (J396.22.w1)

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Authors & Referees

Authors Dr Debra Bourne MA VetMB PhD MRCVS (V.w5)
Referee Susan K. Mikota DVM (V.w72)

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