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BEHAVIOUR - Editorial Comment

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

Home ranges of elephants in Africa vary greatly in size, from about 14 square kilometers to over 3,000 square kilometers. Variations in home range size relate to differences in availability of essential resources such as water and food, but are also affected by factors such as hunting, in that elephants appear to learn where the boundaries of "safe" areas are. Home ranges may also be complex in shape, with long thin corridors linking different areas. Home ranges of family units are relatively stable over long periods, and overlap considerably with one another. Clans also have home ranges and the boundaries of clan home ranges appear to be accepted by clans which are adjacent to one another, although there may be some overlap along the peripheries of the ranges. Bull elephants have their own ranges, occupied by a varying number of bulls; these ranges tend to be larger than ranges of cow elephant clans. Typical daily movements total about seven to eight kilometres (four to five miles); this varies with habitat.

Densities of African elephants may vary from as low as 0.14 elephants per square kilometer to as high as 10 per square kilometer in some areas at certain times of the year. In general, elephants are more concentrated in certain areas during the dry season and more widely dispersed during the wet season. There is no evidence for territoriality. Males disperse from the family group at about the time of puberty.

Elephants may be marked simply, for example with numbers painted on the head and body, or tracked using radio collars; more recently, GPS units have been attached to neck collars.

When a group of elephants is threatened, they typically form a defensive circle, with calves in the centre of the group. The matriarch then investigates the threat.

Elephants are social animals. They communicate with one another using visual, olfactory, tactile, and auditory cues. Elephant society is primarily matriarchal. The basic social unit consists of a female and her offspring (females of all ages and males up to the age of puberty). Family units may join to form larger groups such as "kinship groups" consisting of perhaps four family groups, and even larger clans made up of several kinship groups. Bulls leave or are pushed out of the family group at puberty. While bulls may be found alone they are also found in bull herds, although the composition of a bull group may change constantly. Sometimes males are found with family groups; this may be when a female is in oestrus, but may also indicate males, particularly younger males, maintaining ties with their original family group. Large aggregations of elephants have been described; why they form is not certain. It has been suggested that large aggregations may form in response to threats or droughts, or to exploit a temporary rich food resource, or alternatively that they allow the spilling of large clans and the formation of new groups. Dominance is based on size and strength. Most aggressive interactions are ritualised, such as pushing matches between males in bull herds to determine the place of each individual in the local hierarchy, although fights do occur, particularly between bulls disputing access to a female in oestrus, and one or both parties may be injured or even killed in such encounters. 

Elephants are tolerant towards most other herbivores most of the time, although other species generally defer to elephants. Young elephants may chase other species in play. Elephants may actively chase predators such as lions which are a threat to elephant calves. In montane forests, certain monkeys are found in association with elephants and this may be a mutually beneficial arrangement, with monkeys benefiting from food resources exposed as the elephants fed while the monkeys may warn of the approach of hunters. Various species benefit in drought from water holes dug by elephants. Vegetation may be adversely affected by elephants  but conversely elephants may act as important seed dispersers.

Lions are the main predator on elephant calves; hyenas, crocodiles and wild dogs may also be a threat to calves. There is one hearsay report of lions successfully killing an adult bull elephant.

Elephants do not use nests per se, however they may rest in forest thickets, providing deep shade, in the heat of the day. (B453.9.w9)

Tradition and learning are very important to elephants. They learn information such as choice feeding grounds and the whereabouts of water in the dry season, and, growing up and perhaps becoming matriarchs, pass this information to younger animals. This may enable survival in difficult times. Loss of older animals, particularly matriarchs, as may occur with poaching of the animals with the largest tusks, means that this vital information can be lost.

(References are available in detailed literature reports below)

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Home Ranges and Distances Travelled

Source Information
  • Home ranges have been measured varying from 14 square kilometers to 3,120 square kilometers. This variation depends on the availability of water and the quantity and quality of available food. (B147)
    • In The Lake Manyara area, Tanzania, home ranges of family units and individual bulls were 15 to 52 square kilometers, with wide areas of overlap. (B147)
    • In Tsavo West, Tsavo National Park, Kenya, mean home ranges were recorded as 530 square kilometers; in Tsavo East they were 1,580 square kilometers. (B147)
    • In southern Africa, generally home ranges were small but some elephants were found to move "considerable distances" for short periods. (B147)
  • Ranges vary in size and can be complex in shape, with long corridors linking areas of use. (B285.w3)
    • In Tanzania in a forest areas, ranges have been measured as small as 10 square kilometres ( four square miles), contrasting with 18,000 sq km (7,000 sq miles) in Namibia in a desert area. In Kenya in woodland and brushland, average home ranges of 750 sq km (290 sq miles) were found in an area with abundant food and water, but ranges were 1,600 sq km (617 sq miles) in a drier area. (B285.w3)
  • Typical daily movements total about seven to eight kilometres (four to five miles); this varies with habitat. (B285.w3)
    • In Kenya, only 3.0 km (1.9 miles) were covered by an individual in a forest with good water, compared with 12 km (7.5 miles) for individuals in an arid area. (B285.w3)
  • Elephants show periods of relatively rapid (3.0 - 4.0 km/h (2.0-2.5 mph) movement, in a given direction, generally between two distant sections of their range; this movement, known as "streaking", may be used to traverse dangerous areas between safe areas; it is fairly rare. (B285.w3)
  • Elephants appear to learn which areas are "safe": they will travel to the edge of protected areas then, at the border, turn back. (B285.w3)
  • Boundaries of clan home ranges appear to be accepted by clans which are adjacent to one another, although there may be some overlap along the peripheries of the ranges. (B384.8.w8)
    • In the Kruger Park, clans of breeding herds have discrete home ranges of 126 to nearly 1,000 square kilometres (depending on habitat quality), averaging 450 sq. km. (B384.8.w8)
    • In Tsavo, ranges may be as large as 3,000 sq. km. (B384.8.w8)
  • Home ranges of family units are relatively stable over long periods, and overlap considerably with one another. (B384.8.w8)
    • Home ranges vary in size depending on the habitat (larger in areas of less good habitat) and season (larger in the dry season than in the wet season when food is more plentiful). (B384.8.w8)
    • Home ranges may be reduced due to avoidance of humans. (B384.8.w8)
  • Bull elephants have their own ranges, occupied by a varying number of bulls (e.g. in Kruger Park 10 to 100, average 41 per bull range). (B384.8.w8)
    • Bull elephant ranges are generally larger than ranges of cow clans. (B384.8.w8)
    • Young bulls leaving their family group enter these bull ranges. Initially they may wander widely but most eventually settle in their originally chosen bull range. (B384.8.w8)
    • Members of a bull range generally originate from the closest clan. (B384.8.w8)
    • Young bulls, pre-musth, are most likely to move into new areas. (B384.8.w8)
  • There may exclusive bull areas among elephants, not used by family units. (B396.3.w3)
  • Studies in Lake Manyara National Park, involving 48 family units and 80 bulls, found home ranges of 14 to 52 kmē. (B451.3.w3)
  • In Tsavo National Park, Kenya, ranges of four elephants were measures as 294 to 441 kmē, averaging 350 kmē in Tsavo West, while in Tsavo East (with ecologically poorer habitat for elephant), ranges were found to be 525 - 3,120 kmē, mean 3,120 kmē. (B451.3.w3)
  • In several studies, distances between sightings have been relatively small, e.g. 30 km in one study, 22 km in another, from initial points of darting. (B451.3.w3)
  • Elephants may make short, non-seasonal movements between water sources and food sources; paths may be formed where elephants follow regular routes. (B147)
  • Seasonal movements have been noted in many elephant populations, but are not necessarily true migrations. For example in Rwenzori National Park, Uganda, more elephants were found on the grasslands during the wet than during the dry season, but there were always some in the grasslands and some in the forests. (B451.3.w3)
  • Seasonal movements are related to food availability, which is affected by rainfall. (B451.3.w3)
    • Requirements for other resources, such as sodium, may also affect movements. (B451.3.w3)
  • The longest regular seasonal African elephant migration was recorded as only 30 km [1975 publication]. (B451.3.w3)
    • However, it is possible that longer migrations occurred in the past. (B451.3.w3)
  • Elephants range over larger areas if available food and water is widely dispersed. (B387.w4)
  • Data from five female elephants tracked using GPS radio-collars in the Tarangire National Park, Tanzania, found that three of the elephants mainly used the northern part of the park, but sometimes moved outside to about 10-20 km northeast of the park boundary. The other two, in the southern area of the park, moved about 80 km outside the park boundary. Home ranges were 159, 403 and 660 kmē for the northern elephants and 2104 and 3314 kmēfor the southern elephants. The northern elephants showed considerable home range overlap with one another, as did the two southern elephants with each other. Monthly home ranges varied from 53 to 902 kmē; larger home ranges were used in December or January than in March. The data confirmed the important migration route starting from the northern sector of the Tarangire NP and showed that the northeastern corridor towards Lolkisale Game Controlled Area. The study also showed that elephants within the park had peak movements 8 am to 4 pm while those outside the park showed highest movement rates 4 pm to 12 pm, presumably trying to hide in thicker vegetation during the daytime, avoiding contact with humans. (J398.11.w1)

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Dispersal, Territoriality and Population Densities

Source Information Population Densities
  • African elephant densities have been measured as varying between 0.26 and 5.00 per square kilometer. (B147)
  • It has been calculated that pristine habitats could support about two elephants per square kilometer. (B147) Studies have shown:
    • In the Amboseli National Park, Kenya (about 3,500 square kilometers), an overall density of 0.14 elephants per square kilometer; in the dry season in some areas this increased to 0.4 - 0.9 per sq. km and in some habitats even to 10 per square km. (B147)
  • Often, elephants are concentrated during the dry season/droughts, and more dispersed during the wet season. (B147)
  • A 1979 publication gave the following density estimates, published 1965 - 1975: (B387.w4)
    • Manyara National Park, 5.0 elephants per square kilometre;
    • Arusha National Park, 3.3 elephants per square kilometre;
    • North Bunyoro, 1.9 - 3.8 elephants per square kilometre;
    • Ruwenzori National Park, 2.5 elephants per square kilometre;
    • Ruaha National Park, 1.1 elephants per square kilometre;
    • Kidepo, 0.6 elephants per square kilometre;
    • Tsavo, 0.5 elephants per square kilometre;
    • Mikumi, 0.26 - 0.43 elephants per square kilometre;
    • Luangwa Valley, 2.7 elephants per square kilometre;
    • It was noted that comparison between these estimates was difficult due to co-existence in some areas of resident and mobile elephants for pars of the year. elephants per square kilometre;

Territoriality

  • There are no indications of territoriality in African elephants. (B147)
  • Most studies of elephants have found no evidence of territoriality. (B451.3.w3)
  • One study in Zimbabwe found that groups of large bulls did not stray into one another's territory, and that when such straying did occur, fighting resulted, although small bulls could cross the boundary; these may be group territories and it was speculated that they are held for mating purposes. (B451.3.w3)
  • No evidence was found of territoriality by the Douglas-Hamiltons in the Lake Manyara elephants, nor in another study in western Uganda. (B451.3.w3)

Dispersal

  • Analysis of the age and sex structure of family units in elephant populations in East Africa clearly indicated that males left the family unit at or just after they reached puberty. (P17.21.w1)

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Methods of Marking and Following 

Source Information
  • Most information on the home ranges of elephants has been gathered by radiotracking including, in recent years, use of collars with GPS units. (B285.w3)
  • Some information can be gained by noting positions of marked individuals (e.g. marked with numbers pained on the head and body). More information can be gained by radiotracking, using transmitters attached to neck collars. (B451.2.w2)

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Defensive Behaviour

Source Information
  • When threatened, adults form a defensive circle, facing outwards, with the calves in the middle. The matriarch then checks the threat and may advance on it, spreading ears, trumpeting, growling and even threat charging or actually charging. In confrontations with poachers this means the matriarch is the most likely to be killed first. (B285.w3)
  • To threaten, elephants hold the ears at right angles to the body. (B384.8.w8)
  • When a family of elephants is threatened, the group crowds together. The matriarch identifies the threat and if possible leads the group away rapidly. (B384.8.w8)
    • If one animal is shot, the matriarch will try to assist that animal. (B384.8.w8)
    • If the matriarch is shot, the rest of the group often clusters round the body - and are then easy to shoot. (B384.8.w8)
  • In defensive and aggressive postures (including actual attacks), the ears are held out at right angles to the body, making the elephant appear larger. (B453.1.w1)
  • In the face of possible danger, elephants raise the head and trunk, providing additional height, and extend the ears, increasing apparent breadth. They may make threatening gestures such as scraping the soil with a forefoot, shifting weight between the forefeet or weaving from side to side; the tail may be raised and twisted or wagged. They may give a shrill scream from the trunk. (B453.9.w9)
  • If the intruder is identified and does not retreat, the elephant may mock-charge with the trunk up, stopping after about 30 yards if the intruder retreats, blowing a puff from the trunk before turning and walking back to the other elephants. (B453.9.w9)
  • Sick and injured elephants, as well as those with young calves, or cows accompanying a cow at parturition, are more likely to be aggressive. (B453.9.w9)
  • Elephants which feel trapped, with their exit rout blocked, may be very aggressive. (B453.9.w9)
  • When threatened (e.g. by a sudden noise), the elephants cluster, with small calves under their mother's bellies, larger calves beside or behind their mother, all facing the direction of the sound and testing the wind with their trunks, while the matriarch steps cautiously towards the sound. She may then charge, with trunk down, or mock-charge, with the trunk up. (B453.9.w9)
  • In group defence, the adults may be found on the periphery, facing outwards, with the calves in a cluster in the middle, or the larger elephants pack tightly against the matriarch, with young calves in among their legs and facing away from the source of the action. (B387.w4)
  • If the intruder does not retreat, or is aggressive, a real charge is made, with the head lowered, trunk rolled down and to one side, and the tusks directed at the intruder. (B453.9.w9)
  • An intruder may be grabbed with the trunk and thrown to one side, or may be crushed or stabbed. (B453.9.w9)
  • A serious charge is generally made silently, without preliminaries, although such a charge may also result as an escalation of aggressive posturing. (B387.w4)
  • During a real charge the ears are held back against the body and the trunk is curled up and under. (B384.8.w8)

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Interactions with the same Species

Source Information
  • African elephants are gregarious. (B147)
  • In general, elephants are peaceful. (B147)
  • "The general picture of elephant society that is beginning to emerge is that of a number of bulls loosely associated with a kinship group of females and their young and living within a defined area, which is possibly a group territory. The bulls know each other and keep company but only on a casual basis. There is almost certainly a social hierarchy amongst the bulls but perhaps no one bull is dominant over all the other." (B451.4.w4)

Communication and greeting:

  • Elephants have a well-developed communication system using hearing, vision, touch and chemicals (smell, taste). (J54.19.w8)
    • Acoustic (sound) signals are omnidirectional and can be broadcast widely, including to elephants which are not in sight. This is advantageous for maintaining the cohesion of a group and for locating lost individuals, but less useful for directing at a specific individual. There is a variety of empirical data indicating that African elephants can detect infrasonic calls over distances of at least 4 km. Calls serve a variety of purposes including affiliation (distress and comfort calls), coordination of group activity, group cohesion, agonistic and dominance-related calls, calls indicating emotional states (fear, surprise, social excitement, sexual excitement) and signalling hormonal state (the musth rumble of males and estrous rumble of females). Bulls appear to produce a lower variety of calls than do cows. Calls with a low fundamental frequency and higher harmonics may convey information about the distance of the caller from the recipient, since the low frequency sounds will travel further, therefore at longer distances only these will remain. (J54.19.w8)
    • Visual signals operate over short distances, are quickly degraded by distance or hidden by objects in the environment, and are directional. They are generally less useful therefore for broadcasting a signal widely, but useful for targeting a specific individual, as in aggression or dominance displays, and postural indication of apprehension or submission. Visual displays also can be used to indicate physiological state: the "musth walk" with the bull's head held high and his gait "swaggering", and the female's "oestrous walk" with frequent backwards glances over her shoulder. The urine dribbling of a musth bull may also act as a visual indicator of musth, and the bull's ear wave may provide a visual signal as well as wafting musth scent towards the other elephant. (J54.19.w8)
    • Chemical signals are long-duration signals which are energetically efficient. Sources of chemical signals, such as urine, can accurately indicate the hormonal state of the individual, including oestrus in females and musth in males. They also are used in kin recognition and in particular maternal recognition. Chemical signals such as those associated with musth might be detectable at some distance, although this is not known. (J54.19.w8)
    • Tactile communication is short distance only. Greeting of elephants often involves twinning of trunks and checking one another's mouths with the trunk tip. Checking the mouth may also involve chemical communication, and is also carried out when elephants have not been separated. Adult/calf tactile interactions which may be initiated by the calf or the adult (often not the calf's mother) include greeting/investigating, with the trunk placed around or touching different parts of the other elephant's body, comforting behaviour with the body rubbing against the other elephant, investigating foods (placing the trunk to the mouth, trunk or food of another elephant) and play initiation/invitation - touching or rubbing the other elephant quickly followed by play. Calves also use touch to solicit nursing, pushing against the legs or breasts of their mother or another female (together with a suckle rumble or suckle cry). Touch is also used in agonistic encounters (pushing between bulls) and in discipline - mothers and other elephants slapping calves with the trunk, kicking, poking or shoving. (J54.19.w8)
  • Touch is important to elephants. Elephants frequently touch one another. When greeting one another, they stand close together and entwine trunks. (B285.w3)
  • As part of greeting, elephants put their trunk tips into each other's mouths; this may be a sign of confidence (that the other elephant will not bite off the trunk tip). (B384.3.w3)
  • Placing the trunk tip into another elephant's mouth may be associated with seeking reassurance during stress. (B396.4.w4)
  • Elephants indicate aggression by: 
    • raising the head and trunk;
    • moving the ears so they are extended perpendicular to the body;
    • kicking up dust;
    • swaying of the head;
    • Making a mock charge, or a real charge. (B147)
  • Individuals of a family or body group, when meeting after separation, greet one another with loud rumbles; there are louder and longer lasting if the separation has been longer than those following short separations. (B384.8.w8)
  • Groups of elephants appear to coordinate their movements over long distances (several kilometres); low frequency sounds (infrasounds) are probably used in this coordination. (B384.8.w8)
  • Elephants use deep rumbles as contact calls to keep in touch with one another both while feeding and while moving. (B462.3.w3)
  • Infrasounds involve calls at 14 to 24 Hz, and at e.g. 85 - 95 decibels, some reaching more than 100 dB (measured at five metres). High pressure (volume) infrasounds can carry for considerable distances, even through vegetation, allowing communication at a distance. Infrasonic calls may carry considerable meaning. Studies at Amboseli identified a greeting rumble, contact call, contact answer, "let's go" rumble, musth rumble, female chorus, postcopulatory call and mating pandemonium rumble. (B396.4.w4)
  • The position and movements of the ear are used in signals between elephants. (B384.8.w8)
  • Communication between elephants involves auditory and olfactory cues, also body language and posture. (B387.w4)
  • Methods of communication include:
    • Vocalisations (auditory signals); (B453.9.w9, B396.4.w4)
    • Visual signals, including the position and movements of the ears, head position in relation to the body, tail position, movements of the forefeet and, in particular, movements of the trunk. (B453.9.w9, B396.4.w4)
    • Tactile signals, particularly between dam and calf, and during mating behaviour. (B453.9.w9, B396.4.w4)
    • Olfactory signals, involving secretions from the temporal gland as well as urine and faeces. (B453.9.w9, B396.4.w4)
    • Combined auditory and visual signals, such as slapping the ground with the trunk tip. (B453.9.w9)
  • Communication may be visual, involving ears, trunk, head carriage and even the tail. (B451.3.w3)
  • The trunk is used not just for visual communication but for tactile interaction and for exchange of scents. (B451.3.w3)
    • Trunk-to-trunk and trunk-to-mouth contacts are exchanged by mother and calf, but also by other elephants of all ages and both sexes. (B451.3.w3)
  • Chemical communication appears to be important, although much is still not known. Asian elephants are known to produce a wide variety of chemical signals in urine, on breath, in temporal gland secretions. Reactions to these chemicals indicate that they play roles in male-to-female, female-to-male, male-to male and female-to-female indication of physiological state, also kinship (e.g. elephants recognise the scent of their own mother's urine even after being separated for more than 20 years). (J54.19.w7)

Families and larger units

  • African elephants are generally arranged as a matriarchal society, based around the family unit of a cow elephant and her calves. Small herds exist containing only a single female and her offspring of various ages. More usually, there are groups of about 10 individuals. (B147)
  • Solitary female elephants are very rare; most are part of a family unit. (B147)
  • Two to four family units may join to form a larger "kinship group" or "clan"; it has been noted that such clans of six to 70 individuals are led by a large female. (B147)
    • This female will prevent outsiders from joining the group. (B147)
    • Dominant females normally maintain this position until they die, and are then succeeded by their eldest daughter. (B147)
  • African elephants may be found in groups containing hundreds of individuals or even more than 1,000. It is thought such gatherings are aggregations of smaller herds and individuals, developing in response to disturbances such as drought or human interference. (B147)
  • In Amboseli, Kenya, in the dry season elephant groups rarely exceed 20 individuals, however groups of 400 or more may develop during the rains. (B147
    • It is possible that such aggregations provide collective defence against predators. (B147)
  • Female African elephants live in family groups, with adults being sisters or mother/daughter(s), plus calves. (B285.w3)
  • Young females remain with their natal group. (B285.w3)
  • When the family group gets large, a subgroup of young adults will split off gradually to form their own herd; such related family groups often move in a coordinated fashion. "Kinship groups" or "bond groups" are made up of two to four related family groups which generally travel around together. (B285.w3)
  • The family group is led by the oldest female, known as the matriarch. (B285.w3)
  • If one individual of the group is injured, others often try to assist and help support it to get on its feet and move away with the group. (B285.w3)
  • Young females remain in their family group. (B384.5.w5)
  • In drought conditions, adults will prevent younger elephants from drinking until they have drunk. Younger animals, having been made to wait, may then be left behind as the adults move off to find food. (B384.6.w6)
  • Females will often assist other elephants in difficulty. (B384.8.w8)
  • The family is made up of related animals, such as a female plus one or two offspring, or e.g. five or six related animals, led by the oldest cow. (B384.8.w8)
  • Two or three small family units may be associated, frequently joining to give groups of five, six or even ten animals, then splitting again. These are probably part of an extended family. (B384.8.w8)
  • The matriarch leads the group; she has the greatest knowledge of the best places for feeding and watering in different seasons and rainfall conditions. This knowledge is then passed down through the generations, therefore the home ranges and movement patterns tend to remain the same over long periods. (B384.8.w8)
  • Larger groups of elephants may be noted which share a home range and often come together. (B384.8.w8)
  • Factors which may affect the stability of family units include death of the matriarch, the unit becoming too large, strength of bonds between individuals, physical condition of an important member of the family and attempted immigration of a strange individual. (B384.8.w8)
  • Females may remain with the family group throughout their lives. (B384.8.w8)
  • Large gatherings of elephants may occur. These may be change aggregations at a good food or water source, and show little coordination with one another, or may be very large and coordinated. Gatherings of this second type may form in response to pressures such as loss of matriarchs (shot) and inability to disperse due to human settlements. (B384.8.w8)
  • A "clan" of elephants is made up of a number of related family groups, found within a largely exclusive home range. In the dry season, families of such clan may come together. (B384.8.w8)
  • There is morphological evidence that groups of elephants are composed of related animals. For example, in a study of elephants in East Africa, in the population in Murchison Falls National Park (Uganda) North bank, and excluding solitary animals, 75 (88.2% +/- 7.0%) of those showing reduced numbers of toe nails were found in association with one another and only 10 (11.8%) were not in association with others with this characteristic. Tusk morphology (size and curvature) was also found to be similar within a given family group. (P17.21.w1)
  • In general, the elephant herd is based on a matriarchal pattern with an old female leading a group made up of her daughters plus males up to puberty. Groups may also contain more than one such family units, but groups obviously eventually split up as the group size increased. (P17.21.w1)
  • The basic family unit or sub-clan consists of a female and her offspring (male to the age of sub-adult, and female). These are led by a matriarch, almost always the largest female, who is mother, aunt or older sister to the other adult females, and may have her own young calves. There is usually also a senior herd bull, generally found on the periphery or as far as a mile behind the other elephants, and one or two younger bulls may accompany this male. Old cows usually remain with the herd. While browsing, the herd can be seen to separate into the individual family units. Other groupings include bull herds of up to 12-15 individuals, containing sexually mature males of 15 years and over, and generally led by a senior bull, lone, usually elderly, bulls, which usually do not migrate, and congregations or gatherings of several herds at the end of the rainy season before dispersal in the dry season. (B453.9.w9)
  • Large gatherings, involving hundreds of elephants, have been described. These occurred at the end of the rainy season. Two such gatherings were described in which elephants were found over an area about four or five miles long and 1.5 miles across, in wide swampy pains with high elephant grass, and involved at least 600 and perhaps more than seven hundred elephants. These were describes as being led by a very large bull elephant, followed by three or four other large bulls, then other bulls in groups of two or three further back, while on either side were family herds as well as groups of three or four younger bulls. It was speculated that such gatherings allowed the formation of new herds, with large clans splitting up and "up-and-coming" bulls attaching themselves to new clans. (B453.8.w8)
  • In the event of the death of the herd matriarch (e.g. when she is shot), the next largest cow assumes the position of leader. This is possibly a sister or the eldest daughter of the previous matriarch; it is not known whether this female is already recognised as "herd deputy". (B453.9.w9)
  • At Manyara, it was noted that stable family units, averaging about ten elephants per group, belonged to a larger, related, kinship group. Family units within such a kinship group tended to remain close to one another and sometimes came together to form a larger group. (B462.4.w4)
  • If one elephant was darted, other elephants in the same family unit, or sometimes even from other family units, sometimes stayed with the darted individual, protected it and tried to raise it back to its feet. This behaviour when a calf was darted was such that it was impossible to approach the immobilised animal. (B462.7.w7)
  • Calves of a dead female, or other adults, may stay with a dead elephant for some time. (B462.16.w16)
  • Elephant society is primarily matriarchal and the basic social unit consists of a female and her immature offspring. As daughters mature and bear calves, a larger matriarchal family is formed. (B451.3.w3)
    • Groups with more than one female of about the same age, and no obvious single matriarch, may represent a family which has lost its matriarch; the females might gradually split up into small units, each with their offspring. (B451.3.w3)
    • Groups with more than one matriarch are also found, possibly formed from several family groups coming together temporarily. (B451.3.w3)
    • If an animal in a group is darted, the matriarch will guard the darted individual. If the matriarch is darted, the group first runs to her (which is used in whole-family culling operation) then they appear unsure what to do and can be pushed away; the group will then stand and watch. (B451.3.w3)
    • In some areas, family units appear to be less cohesive, sometimes, such as in the dry season or after a calf is born, splitting into sub-units, but then coming together again. (B451.3.w3)
    • Large groups may represent a response to threat and/or to loss of leaders. (B451.3.w3)
    • Observations of family units accompanied by sexually mature males, indicates that while in some cases the male may be present because a female is in oestrus, this is not always the case. It is suggested that young bulls may retain contact with their family units: the average age of the bulls accompanying families in this way is 24 years, compared with an average age of 31 years for bulls in bull groups. (B451.3.w3)
    • The mean size of a basic family unit appears to be about 3 (average 2.8). Different studies have found family group sizes averaging 7.8 to 12.3 in different areas in one study, and 10 in another study. (B451.3.w3)
    • There appear to be higher, larger levels of elephant organisation, with some groups, probably related, coming into contact more often than would be expected by chance. (B451.3.w3)
      • A "kinship group" was defined by Douglas-Hamilton as being an association of three of four family units, totalling up to 50 elephants; units within such a kinship group in Lake Manyara National Park were noted to keep close together for long periods, but also to split up by considerable distances. (B451.3.w3)
      • Within a kinship group, the oldest matriarch is dominant and this appears never to be challenged. (B451.3.w3)
      • Even larger associations, clans, may be formed from several such kinship groups. Such a kinship group consists of females and prepubertal males, and the animals within it share a common home range. They may sometimes come together, forming herds of more than 100 elephants. (B451.3.w3)
  • Large temporary gatherings may be to discourage predator harassment, or to exploit a rich, but short-lived source of food. (B387.w4)
    • "Seasonal movements, dispersals and congregations are therefore likely to be the result of complex interaction in which nutritional and physiological needs combine with strategies for security; to this should be added the element of tradition that may guide particular populations along paths learnt from the older elephants." (B387.w4)
  • Clumping of elephants, with formation of relatively large group sizes was noted on the periphery of Kabelega (Murchison Falls) National Park, compared with group numbers in the centre of the park. It was suggested that this indicated "that a breakdown or change in the behavioural mechanisms promoting dispersal was involved." (B387.w4)
  • Populations of elephants have been described as forming six hierarchical tiers of organisation, with tier-one units being the mother and calf, tier two the family, tier three the bond group or kinship group, tier four the clan, tier five a subpopulation and tier six a population. Cluster analysis was used to quantitatively identify the existence of four social tiers in African elephants. The second-tier units or families remained stable across seasons, and the number of these units increased as the population size increased. Second-tier units led by older matriarchs, 35 years or older, and likely to be grandmothers, were larger than those led by younger matriarchs. Aggregation of elephants into larger, third- and fourth-tier units occurred less frequently and were significantly affected by season; there appeared to be a trade-off between the social and ecological benefits of formation of larger units and the ecological costs (for example from resource competition). Additionally, the reproductive state of females, and the age and development of their calves, are likely to affect the stability of social units. (J334.69.w1)
  • Elephants coordinate their associations and movements, sometimes over long distances - up to e.g. 5 km; this might be carried out using very low frequency sounds (infrasound). (J396.22.w1)
  • Elephants from a given family or bond group, which have been apart for several hours, use a "greeting rumble" when they are reunited; if they have been apart for a longer period then the greetings are more intense: as well as loud rumbles they trumpet, scream, urinate, defecate and contact one another physically. (J396.22.w1)
  • An elephant which is apparently trying to locate its family may use contact calls and contact answers for several hours. (J396.22.w1)
Bull elephants
  • Solitary bull elephants are common, although males may form temporary associations, which individuals may join and leave at intervals; these bachelor herds are usually small, although herds containing as many as 144 individuals have been recorded. Elderly bulls are often solitary. (B147)
  • When young males reach puberty, at eight to 20 years old, the older females drive them out of the family group; the young bulls then join or form their own bachelor groups. (B147)
  • Adult males join the female-based family groups only when there is a female in oestrus. There is a bull in association with any given family group about 50% of the time. (B147)
  • Males compete with one another for dominance within their groups. (B147)
  • Males compete with one another for access to females in oestrus. (B147)
  • Dominance struggles between males are usually resolved by pushing and light tusking. ((B147))
  • Battles for access to females in oestrus may be more serious and sometimes involve fatal injuries from the tusks. (B147)
  • Young males either leave or are pushed out of their natal group at puberty. (B285.w3)
  • Adult bulls spend some time alone, and much time in small male groups. These groups change in composition and size but recent research in Kenya has indicated that the same individuals may form associations repeatedly. (B285.w3)
  • Young males are driven away from the family, by their own mothers or by a bull when the mother comes into oestrus. The young bull may be driven away at about 15 years of age: at Amboseli it occurred at nine to 19 years old, on average at 14 years. (B384.5.w5)
  • Males gradually become independent at about 14 years old, perhaps making a group with other young bulls. (B384.8.w8)
  • Bull elephants do not appear to have particular attachments, but nevertheless are generally relatively close to one another. Young bulls may form small bachelor groups or may join with an older bull. Some bulls are solitary. (B384.8.w8)
  • A radio-collared male followed for three weeks at Manyara showed no attachment to any particular individual or group. He was alone some of the time, but never for a whole day, but also spent time with 12 bulls, some larger and some smaller, and temporarily associated with four different family units. The longest time spent with another bull was five days. (B462.7.w7)
  • A young bull (about 19 years of age) was noted to have serious fights only with bulls of his own age class, sometimes resulting in deep gashes across the trunk, while relationships with older bulls were friendly and he never challenged them over e.g. access to a waterhole. (B462.14.w14)
  • Bulls continue to play-fight for their whole lives, which probably provides a means whereby they can check their position in the social hierarchy. Generally, adult bulls appeared to have a size-based hierarchy and conflicts were resolved using mild threat gestures, with serious fights being rare. (B462.14.w14)
  • Bulls generally show gentle behaviour towards females and calves, although aggression has been seen. (B462)
  • 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)
  • Bulls are independent but are sociable and are usually found within a mile of either another bull or a family unit. A bull herd of 14 was seen but was considered unusually long and this number of bulls did not remain together for very long. (B462.14.w14)
  • Bulls formed brief associations with females when the females were in oestrus. (B462.14.w14)
  • A 19-year-old bull at Manyara was noted to still be attached to a family unit, usually trailing it by several hundred yards. Eventually, having tended to linger behind more and more when the family moved, one day he turned off in the opposite direction and walked out of the normal range of his kinship group. (B462.8.w8)
  • Within a bull range the individuals develop a dominance hierarchy. (B384.8.w8)
    • Young bulls challenge older bulls to learn their place in the hierarchy; large, dominant bulls do not challenge subadult bulls. (B384.8.w8)
    • Musth affects the hierarchy: normally, the older, larger bull wins contests, but a bull in musth usually wins encounters with a non-musth bull, even when the non-musth bull is larger. (B384.8.w8)
    • Non-musth bulls generally avoid musth bulls, and musth bulls generally avoid one another. (B384.8.w8)
    • Musth bulls generally fight in a ritualised manner, with a forehead-to-forehead approach and clashing of tusks; this is still dangerous and can be fatal. (B384.8.w8)
    • Fights are not common, but when they occur they can be long lasting (up to six hours) and may result in death of one elephant, breaking of tusks and other injuries. The victor chases the loser. (B384.8.w8)
  • After leaving the family herd at puberty, young males may spend a variable amount of time as solitary animals before joining a bull herd. (P17.21.w1)
  • At puberty, males generally join a bull herd. (P17.21.w1)
  • Rank among bulls may be determined by fighting, or without fighting, the smaller, weaker bull withdrawing or assuming a submissive posture. (B453.9.w9)
  • Bulls reaching sexual maturity may leave the family herd of their own choice, or may be chivied by the females. (B451.3.w3)
    • Having left the family unit, bulls soon join bull groups. (B451.3.w3)
    • In 110 bull groups culled in Uganda, group size ranged from one to 11, averaging 3.1 south of the Nile River and 2.4 north of the Nile. (B451.3.w3)
    • An aerial survey of Tsavo National Park, Kenya, gave bull group sizes of one to 14, average 2.4, for 427 groups. (B451.3.w3)
    • Members of bull groups have only loose attachments to one another, with individuals leaving and joining. (B451.3.w3)
    • One function of bull groups may be to allow elephants to sort out a dominance order. (B451.3.w3)
    • Bulls may associate partly simply due to a desire for company. (B451.3.w3)
    • Bull groups may contain males of all ages. (B451.3.w3)
    • Young bulls may benefit from association with an old bull. (B451.3.w3)
Dominance, hierarchy and fighting:
  • Hierarchy appears to be governed by size, strength and age. (B453.9.w9)
  • As calves and juveniles, elephants of both sexes play-fight. (B451.3.w3)
  • In very young elephants, of both sexes to the age of puberty, challenges involve head nodding, shaking and crashing through the grass, then pushing contests or head-on butting. At puberty this behaviour stops in females but males become more aggressive. (B387.w4)
  • Aggression is generally ritualised, involving mainly intimidation displays, reducing the chance of injury as well as conserving both energy and time. (B387.w4)
    • Elephants try to gain a height advantage during conflicts, including using termite mounds or the slope of a hillside. They get their heads as high as possible and raise the head, with ears spread wide, giving the appearance of a large single surface. (B387.w4)
    • Prior to raising the head, the elephant may smash bushes and make upward scoops with the trunk at the same time as producing blasts of noise through both trunk and mouth. (B387.w4)
    • Displays may include a series of short advances, with a shallow scoop of the tusks then standing high with the head and ears fully elevated. (B387.w4)
  • Dominant individuals direct foot scuffs or backward kicks towards a sparring partner which is much smaller. (B387.w4)
  • Tossing vegetation or other objects, and tearing up and smashing vegetation may be displays directed against an intruder who does not leave in response to threats. (B387.w4)
  • Young elephants with tusks not yet grown push against one another with the trunk basses touching and trunks intertwined. Older animals may clash foreheads but more usually lock heads together with the tusks and trunks, then try to throw the opponent off balance using twists directed down and sideways. (B387.w4)
  • A submissive or beaten elephant flees or presents its rump to the dominant animal. The dominant animal may then mount the submissive elephant. (B387.w4)
  • An elephant may indicate submission during a fight by placing its trunk tip in the other elephant's mouth. (B387.w4)
  • Postures of the dominant and inferior elephant differ: head up, extended and flapping ears, and sometimes trunk brought forward to touch the side of the other elephant's head by the dominant animal, compared with lower head carriage, ears drawn back and trunk tightly curled in the inferior elephant. (B387.w4)
  • Most fights between bulls are pushing matches to determine position in the dominance hierarchy, however serious fights, using tusks, do occur and bulls are sometimes killed in such fights. (B451.3.w3)
    • An elephant losing in a dominance contest will break away. (B451.3.w3)
    • Serious injuries and deaths are probably caused by closely matched bulls disputing rights to an oestrus female. (B451.3.w3)

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Interactions between Species

Source Information
  • Females with young "are unpredictable". (B147)
  • Calves may chase other species such as warthog and waterbuck. (B385.5.w5)
  • Elephants may be seen with egrets, oxpeckers and piapiacs on their backs; these birds presumably peck at insects on the elephant. (B453.8.w8)
  • In montane forests, monkeys such as Sykes' monkey (Cercopithecus mitis - Diademed monkey) and vervet monkeys (Chlorocebus aethiops - Savanna monkey) may be found associated with elephants. The monkeys may benefit from exposure of foods as the elephants browse (e.g. insect larvae under bark); the elephants may benefit from fruits dislodged by the monkeys from high branches. Additionally, monkeys may act sound an alarm as a hunter approaches. (B453.8.w8)
  • Warthogs have been seen associated with elephants, rooting around as the elephant digs roots and tubers. (B453.8.w8)
  • Various species benefit from water holes dug in dry river beds by elephants. (B453.8.w8)
  • The elephant is tolerant towards most species "but not towards the black rhinoceros which formerly shared most elephant habitats as a competitor, nor towards the ratel or honey badger, zorilla and driver ants." (B453.8.w8)
  • In general, elephants tolerate other species and take little notice of them (except for predators when the elephants have young calves). (B453.9.w9)
  • Elephants generally tolerate other herbivores, although other species generally get out of the way of elephants. (B451.3.w3)
    • Incidents are known of other species defying elephants, for example a bull buffalo standing up to an elephant at a salt lick. (B451.3.w3)
    • A group of elephants killed a rhinoceros which had attacked an elephant calf. (B451.3.w3)
    • Hippos and elephants appear to be mutually tolerant, but hippos generally move out of the way of elephants. (B451.3.w3)
    • Young elephants may charge smaller species such as antelope, zebras and warthogs, apparently in play. (B451.3.w3)
    • The supposed animosity between black rhinoceroses and elephants "is rarely in evidence." (B451.3.w3)
  • African elephants may chase predators such as lions. (B451.3.w3)
  • Elephants will generally use an intimidation display against a competitor or predator species. (B387.w4)
  • Elephants may have a considerable impact on the vegetation, with for example preferred forage trees being reduced in number and disliked or distasteful species increasing. (B387.w4)
  • Elephants may act as important seed dispersers; seeds are commonly seen germinating in old piles of elephant dung. (B387.w4)

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Predation in the Wild

Source Information
  • Young elephants may be predated by lions, hyenas or crocodiles. (B285.w3)
  • Lions are the only predator of elephants (other than humans), taking calves and even young elephants up to 12 years old. (B384.5.w5)
  • Young elephants are at risk of predation by lions. (B451.3.w3)
  • Calves may be predated by lions, hyenas, crocodiles and wild dogs. (B453.7.w7)
  • There is a hearsay report of lions successfully killing an adult bull elephant. (B451.3.w3, B453.8.w8, B387.w4)
  • A calf of seven or eight years old was found to have lost part of its trunk and been wounded deeply on its legs, shoulders and neck during the struggle while being killed by lions. (B387.w4)
  • Calves are sometimes predated by lions. (B462.16.w16)
  • In the Gounda study area in the centre of the Gounda-St Floris National Park, Central African Republic, one of the resident lion prides was observed with nine elephant prey over the period 1981-1984. One elephant killed was a young bull, about 10 years old, weakened by several spear wounds about six months earlier; he was killed by bites to the throat. The others were all under seven years of age. Two of the juveniles in poor condition wandering alone were seen with deep scratches on their flanks indicating unsuccessful lion attacks. It was thought that this pride had taken to preying on juveniles left alone due to poaching. It was noted also that in Chobe National Park, Botswana, during an extended drought in 1987, two separate pairs of pride males were considered experts in killing elephant calves. Orphaned calves quickly lose condition and are probably relatively easy prey for a lion pride. (J183.29.w1)
  • A study of a lion pride in the Chobe National Park, Botswana, near a waterhole found that the lions preyed on elephants, killing 74 elephants over a period of four year. One was a bull that had been injured by another bull elephant. All the others were elephants from breeding herds of females and their young. Predation was rarely attempted on calves under four years of age, which remained close to their mothers. Two-thirds of those killed were four to 15 years of age, usually individuals on the periphery of, or straggling behind, the herd; most were killed more than 50 m from the herd. The lions appeared to be increasing their elephant hunting success: 11 in 1993, 17 in 1994, 19 in 1995, 27 in 1996. It was noted that there were reports of lions killing elephants elsewhere in Botswana at waterholes, along rivers etc. (J183.44.w1)

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Nest Use

Source Information
  • Elephants may rest in forest thickets, providing deep shade, in the heat of the day. (B453.9.w9)

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Intelligence and Learning

Source Information
  • A number of factors favour the development of intelligence in elephants:
    • "Elephants need to differentiate identities, to record memories of other elephants' behaviour, and to store experiences of droughts, dangerous places and situations, and promising feeding sites." (B285.w3)
    • "Given their long lives, there is a survival premium for families led by matriarchs that make the right decisions on movements in times of peril or drought." (B285.w3)
  • Tradition and learning are very important to elephants. They learn information such as choice feeding grounds and the whereabouts of water in the dry season, and, growing up and perhaps becoming matriarchs, pass this information to younger animals. This may enable survival in difficult times. (B451.3.w3)
    • Loss of older females to poachers reduces the amount of local tradition which is passed on to he younger elephants. (B451.3.w3)
  • The cultural knowledge of the elephant's environment is passed down between the generations and may include information such as where to find water in drought which may occur only at interval of decades. (B389.21.w21)
  • Calves learn which food to eat by sampling food types they see eaten by other members of the herd. (B285.w3)
  • Elephants pass on their experience from one generation to the next and can perpetuate "traditions" within their family units. (B462.17.w17)

Information on tool use is provided in African Elephant Loxodonta africana - Activity Patterns, Grooming and Navigation Behaviour (Literature Reports) - Activity Patterns

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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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