Animal Info - African Elephant

(Other Names: 非洲象, ελέφαντας, アフリカゾウ, 아프리카 코끼리,  Африканский Слон, Afrikanischer Steppenelefant, Dôli, Elefante Africano, Eléphant Africain, Eléphant d'Afrique, Elkanjavwini, Elu, Enyi, Erin, Fil, Giwa, Indlovu, Indyamba, Inzovu, k, Khoab, Llou, L'xo, Marodi, Mbala, Mbaus, Ndhlopfw, Ndhlovu, Ndlovu, Ndovu, Niay, Njogu, Nyiiwa, Nzou, O Lenkaina, Olifant, Oltome, Ondjamba, Ondjou, Qgo, Swah, Tembo, Tlou, Tou, Xóà, Zehon)

Loxodonta africana* 

(*See Taxonomy)

Status: Vulnerable


Contents

1. Profile (Picture)
2. Tidbits (Elephants and People)
3. Status and Trends (IUCN Categories, Countries Where Currently Found, Taxonomy, Population Estimates, History of Distribution, Threats and Reasons for Decline)
4. Data on Biology and Ecology (Size and Weight, Habitat, Age to Maturity, Gestation Period, Birth Season, Birth Rate and Population Growth, Early Development, Dispersal, Maximum Reproductive Age, Maximum Age, Diet, Physical and Behavioral Characteristics (The Trunk, The Tusks, The Teeth, The Ears, Communication by Sound, Activity Patterns, Bathing, Sleeping, Movement, Migration, Reproductive Flexibility, Elephant Graveyards), Social Organization (The Family, Relations between Family Members, Larger Groups, The Greeting Ceremony, Aggression), Gender Distribution, Mortality and Survival, Density and Home Range)
5. References


Profile

Pictures: African Elephant #1 (37 Kb JPEG) (Nature-Wildlife); African Elephant #2 (Small Family) (56 Kb JPEG) (WWF); African Elephant #3 (Large Family Drinking) (63 Kb JPEG) (CITES); African Elephant #4 (Greeting Ceremony) (74 Kb JPEG) (Naturefoto-Online); African Elephant #5 (Mother and Baby) (113 Kb JPEG) (Czech Web Site)

Male African elephants average about 3.2 m (10.5') in height and weigh an average of 5000 kg (11,000 lb).  Females are smaller. The African elephant occurs in a wide range of habitats, including dense forest, savanna, grassland, marshes, thornbush, semidesert scrub and arid desert. It feeds on a wide variety of plants, ranging from grass to the leaves, twigs and flowers of trees. The forest elephant also feeds on a wide variety of tree fruits. While searching for food and water, the African elephant can walk up to 500 km (300 mi). It can run (for short distances) as fast as 40 kph (25 mph), and it is an excellent swimmer.

The major unit of elephant society is a stable, matriarchal family of cows and their calves. A typical family includes an average of 10 individuals, usually consisting of an older female, her dependent offspring, and her adult daughters with their immature offspring. Males are usually driven out of the family when they attain puberty, after which they remain solitary or temporarily congregate into bachelor herds. African elephant families are not territorial. In fact, they appear to have a social affinity for one another and can be found in larger groups of various sizes. 

Communication within a family takes many forms, including vocalizations, frequent touching with their trunks, and body postures. Family members are strongly bonded with one another. The strength of the bonds is demonstrated by the greeting ceremony. If two subgroups of a family have been separated for a few days, the greeting ceremony accompanying their reunion will include rumbling and trumpeting, clicking their tusks together, entwining their trunks, flapping their ears, and generally showing great excitement.

In the past, the African elephant occurred throughout Africa, from the Mediterranean Sea to the Cape of Good Hope at the bottom of the continent, except in parts of the Sahara and some other desert regions. It disappeared from north of the Sahara by about the 6th century AD. The elephant is still relatively widely distributed south of the Sahara, but populations are now fragmented, especially in West and East Africa. Elephant population trends during the 20th century differed by region. In East Africa, there was a population maximum around 1970. In southern Africa, elephant numbers were at their lowest around the beginning of the 20th century and have been increasing steadily since then. In West Africa, major declines probably occurred well before the turn of the 20th century, and the population has remained at low levels ever since. 

The decline of the African elephant can be linked to three major factors: 1) desertification, which was a major cause for the disappearance of the species in North Africa and the Sahara; 2) the killing of elephants to satisfy the demand for ivory, which has been the major factor in reducing elephant populations throughout most of history; and 3) conflicts between elephants and humans for the use of land. Although the last factor may have been of minor significance until recently, the recent rapid growth in human populations in Africa has resulted in large areas of the continent now being permanently unsuitable for elephants. 


Tidbits

*** The African elephant is the largest living land animal. The largest known specimen, a bull elephant, died in 1955. It weighed 10,000 kg (22,000 lb) and stood 4 m (13') at the shoulder. (Brookfield Zoo) 

*** Elephants are highly intelligent and have been observed both using and manufacturing tools (Laursen & Bekoff 1978).

*** "Elephants seem to have some concept of death. Unlike other animals, elephants recognize one of their own carcasses or skeletons. When they come upon an elephant carcass they stop and become quiet and yet tense in a different way from anything I have seen in other situations. First they reach their trunks toward the body to smell it, and then they approach slowly and cautiously and begin to touch the bones, sometimes lifting them and turning them with their feet and trunks. They seem particularly interested in the head and tusks. They run their trunk tips along the tusks and lower jaw and feel in all the crevices and hollows in the skull..." (Moss 1988)

*** Just as humans are right-handed or left-handed, elephants are right-tusked or left-tusked. The dominant tusk is rounded and more worn at the tip. (Ricciuti 1993)

*** The ears of elephants can be used to identify individuals, using the unique vein patterns in the ears and the patterns of holes, nicks and tears on the edges of the ears (Moss 1988)

*** A number of characteristics distinguish the African elephant from the Asian elephant (Elephas maximus). The African elephant is taller and generally weighs more. It has larger ears, its highest body point is the shoulder, there are two fingerlike "processes" at the tip of the trunk, and both sexes carry tusks and are swaybacked. In contrast, the Asian elephant has smaller ears, its highest body point is the head, there is only one finger-like "process" at the trunk tip, only males generally carry tusks, and the back is level or convex. (Laursen & Bekoff 1978)

*** Elephants and People:

The first recorded observations of African elephants made by man are rock etchings. A number of them are found in the Sahara which, in the period of 5000 - 11,000 BC, was not a desert, but bore vegetation that made it a suitable habitat for elephants. (Douglas-Hamilton & Douglas-Hamilton 1975)

"Some portions of the elephant are, however, grateful even to European palates, and the foot, when baked, is really delicious..." (Wood 1860)

The African elephant is intelligent and not difficult to tame, but it has not been utilized by people for labor and transportation to the same extent as the Asian elephant. Domestication of the African elephant for military purposes began in 285 BC with Ptolemy II along the Red Sea and later at Adulis in Ethiopia. Hannibal used African elephants from the Atlas Mountains against the Romans [as described below]. In 1900, King Leopold II of Belgium established a training station for elephants in The Democratic Republic of the Congo, which still operates at Gangala na Bodio on a small scale for local work.  A few African elephants have been employed in other countries to carry tourists on safari.  The training of elephants of ages 12 - 15 years takes 10 - 12 months. (Laursen & Bekoff 1978, Ricciuti 1993, Nowak 1999) 

In 218 BC Hannibal set out to attack the Romans in Italy. He marched through Spain and southern France and headed for the Alps with an army that included 37 African war elephants. The elephants were of great help to him in the mountains since his enemies were afraid to approach the unusual animals. The elephants also proved to be excellent climbers. In the autumn of 218 BC, Hannibal reached the Po valley in Italy with all 37 elephants having survived the crossing of the Alps. They were again of help to him during the initial confrontation with the enemy. A fearless attack by the elephants completed the total defeat of the enemy forces. However, the rigors of the strenuous mountain crossing and the winter snow and cold finally were too much for the elephants, who were native to the warm region of North Africa, and they gradually succumbed. (Groning and Saller 1998)

The African elephant does not ordinarily threaten people, although conflicts occur, especially in areas of expanding agricultural development. In Kenya, 108 persons were killed by elephants from 1990 - 1993. In Zimbabwe, elephants killed 500 persons between 1982 and 1989. (Nowak 1999) 

The sport hunting of elephants is permitted under the legislation of a number of countries, and the following countries have CITES export quotas for elephant trophies: Botswana, Cameroon, Gabon, Mozambique, Namibia, South Africa, Tanzania, Zimbabwe (IUCN 2004).


Status and Trends

IUCN Categories

Note: It is not possible to state whether the change in 2004 to a rating of "Vulnerable" is due to real changes in the status of the African elephant, to the availability of better or more complete information, and/or to the use of different methods of assessing the elephant population (IUCN 2004).

Countries Where the African Elephant Is Currently Found:

2004: Occurs in Angola, Benin, Botswana, Burkina Faso, Cameroon, Central African Republic, Chad, Congo, The Democratic Republic of the Congo, Côte d'Ivoire, Equatorial Guinea, Eritrea, Ethiopia, Gabon, Ghana, Guinea, Guinea-Bissau, Kenya, Liberia, Malawi, Mali, Mozambique, Namibia, Niger, Nigeria, Rwanda, Senegal, Sierra Leone, Somalia, South Africa, Sudan, Swaziland (re-introduced), Tanzania, Togo, Uganda, Zambia, and Zimbabwe. Extinct in Burundi, The Gambia, and Mauritania(IUCN 2004)

Taxonomy:

Preliminary genetic evidence suggests that there may be at least two species of African elephants; namely, the savanna elephant (Loxodonta africana) and the forest elephant (Loxodonta cyclotis). These two groups have previously been treated as subspecies, Loxodonta africana africana and L. a. cyclotis. The possibility of a third species of African elephant, the "West African elephant" (inhabiting both forests and savannas in West Africa), has also been raised (Eggert et al. 2002). However, the African Elephant Specialist Group of the IUCN believes that premature allocation into more than one species of African elephant may leave forest/savanna elephant hybrids with an uncertain conservation status. For this reason, they decided to conduct the 2004 Red List assessment by treating all African elephants as a single species (IUCN 2004)

The savanna elephant differs from the forest elephant in having a larger body size than the forest elephant, sparser hair covering, triangular-shaped ears rather than smaller round ears, gray skin rather than brown as in the forest elephant, and horizontal, thick, curved tusks as opposed to the straight, slender downward-pointing tusks of the forest elephant. 

Population Estimates:

[Note: This is a long section. You can skip to the next section if you wish.]

[Note: Figures given are for wild populations only.]

Notes: 
  • *Because of the wide range in the type and accuracy of surveys of African elephant populations in different countries, recent continent-wide estimates for the African Elephant Database listed above (Said et al. 1995, Barnes et al. 1999, Blanc et al. 2003) include four different categories of estimates: "Definite," "Probable," "Possible," and "Speculative." For population estimates given here where a range of two numbers is listed followed by four numbers in parentheses, the first number in the range is the (rounded off) "Definite" estimate and the second number is the (rounded off) sum of the "Definite," "Probable," "Possible," and "Speculative" estimates. The four numbers in parentheses list the "Definite," "Probable," "Possible," and "Speculative" estimates in that order. 
  • There are no credible estimates for the worldwide (i.e. continental) African elephant population prior to the late 1970s (IUCN 2004). 
  • **The Definite, Probable, Possible, and Speculative figures reported at the continental level for 2002 are each higher than in the previous report for 1998. While these changes suggest an increase in the continent-wide elephant population between 1998 and 2002, their interpretation is complicated and may be confounded by a large number of factors. Consequently, these data cannot give an indication of overall changes in the continent-wide elephant population in the period between the two reports. (Blanc et al. 2003) 

History of Distribution:

In the past, the African elephant occurred throughout Africa, from the Mediterranean Sea to the Cape of Good Hope at the bottom of the continent, except in parts of the Sahara and some other desert regions. It disappeared from north of the Sahara by about the 6th century AD. The elephant is still relatively widely distributed south of the Sahara, but populations are now fragmented, especially in West and East Africa.  Elephant population trends during the 20th century differed by region. In East Africa, there was a regional population maximum around the late 1960s and early 1970s. In southern Africa, elephant numbers were at their lowest around the beginning of the 20th century and have been increasing steadily since then. In West Africa, major declines probably occurred well before the turn of the 20th century, and the population has remained at low levels ever since. There is insufficient information on sub-regional trends in Central Africa prior to 1977, but elephant populations have probably declined since. Overall there remains insufficient information to determine a current trend at the continental level.  (Sharp 1997b, Nowak 1999, IUCN 2004)

Elephants now occupy around 5.3 million sq km (2.0 million sq mi) of their former range of 30 million sq km (11.5 million sq mi) (Leader-Williams 1996, Blanc et al. 2003).

In areas of Africa where forests and savannas come together, elephants that are hybrids of the savanna and forest elephants are found. It is now known that this area of hybridization is very extensive. (Cumming et al. 1990) 

There are still a few small populations of desert elephants, such as those in northwest Namibia and an isolated herd in the Gourma region of Mali. The Gourma (Mali) population is the northernmost viable population remaining in Africa. (Stuart & Stuart 1996)  

Distribution Map #1 (37 Kb GIF) (WWF 1997)
Distribution Map #2 (6 Kb GIF) (Afr. Ele. Database 2001)
Distribution Map #3 (Afr. Ele. Status Rpt. 2002) [Note: This is an excellent, detailed map.  However, accessing it requires first downloading a 308 page pdf file (using Adobe Acrobat Reader) before the desired map is exhibited.  This takes a long time if you don't have a fast Internet connection.]

Threats and Reasons for Decline:

The decline of the African elephant can be linked to three major factors: 1) desertification, which was a major cause for the disappearance of the species in North Africa and the Sahara; 2) the killing of elephants to satisfy the demand for ivory, which has been the major factor in reducing elephant populations throughout most of history; and 3) conflicts between elephants and humans for the use of land. Although the last factor may have been of minor significance until recently, the recent rapid growth in human populations in Africa has resulted in large areas of the continent now being permanently unsuitable for elephants. (Cumming et al. 1990, Kingdon 1997, Burnie & Wilson 2001, IUCN 2004) 

Other factors that have contributed to the African elephant’s decline include poverty; civil unrest; the easy availability of firearms; antagonism to wildlife (especially elephants causing crop damage); undermanned, underfinanced and undertrained wildlife authorities; lack of liaison between ivory traders and conservation authorities; and lack of understanding of the seriousness of the problem. (Cumming et al. 1990) 

From prehistoric times to the present day, ivory has been sought by man as a luxury and a tool. The Trojans wore buckles and pins fashioned from the elephant's tusks, and adorned their war chariots with bits of ivory. King Solomon sat on a throne of ivory as he delivered his judgments. The Greeks cut ivory into statues of their gods; the Romans honored illustrious men with handsomely chiseled writing tablets and scepters carved in ivory. From 1860 - 1930, 25,000 - 100,000 elephants were killed annually for their ivory, mostly to supply material for the manufacture of piano keys in Europe and the United States.  In 1990, the international sale of ivory was banned, but demand remains. Despite concerns about poaching, controlled sales resumed in 1997 in three African countries (Botswana, Namibia and South Africa), and some of the revenue is used for conservation purposes. Poaching for ivory has lessened from its 1970s’ and 1980s’ peaks but is still substantial in some localities.  (La Monte & Welch 1934, Kingdon 1997, Nowak 1999, Burnie & Wilson 2001)


Data on Biology and Ecology

Size and Weight:

Savanna Elephant:

Shoulder height: Female: 2.4 - 3.4 m (avg 2.5 m) (7.9 - 11.2' (avg 8.2')); Male: 3 - 4 m (avg. 3.2 m) (9.8 - 13.1' (avg 10.5')) 
Weight: Female: 2400 - 3500 kg (avg 2800 kg) (5280 - 7700 lb (avg 6160 lb)); Male: 4000 - 6300 kg (avg 5000 kg) (8800 - 13,860 lb (avg 11,000 lb))

Forest Elephant

Shoulder height: Female: 1.6 - 2.4 m (avg 2.1 m) (5.2 - 7.9' (avg 6.9')); Male: 1.6 - 2.9 m (avg 2.5 m) (5.2 - 9.5' (avg 8.2'))
Weight: Female: 1800 - 2500 kg (3960 - 5500 lb); Male: 2800 - 3200 kg (6160 - 7040 lb) 

(Stuart & Stuart 1996, Nowak 1999)

The largest known African elephant, a savanna male shot in Angola in 1955, weighed 10,000 kg (22,000 lb) and measured 4 m (13.1') at the shoulder (Macdonald 2001).

Habitat:

The African elephant occurs in a wide range of habitats, including dense forest, open and closed savanna, grassland, marshes, thornbush, semidesert scrub and arid desert. The forest elephant is found in the tropical rainforest zone of West and Central Africa, while the savanna elephant occurs in the remainder of the range. The African elephant is found over a wide altitudinal range – from ocean beaches at sea level up to mountain slopes at 5000 m (16,400') above sea level. It ranges from the northern tropics to the southern temperate zone (approximately between 16.5° North and 34° South latitude). (Nowak 1999, IUCN 2004)

The African elephant’s main habitat requirements are: plentiful food in the form of grass or browse, some shade, and a supply of fresh water (although its ability to forage as far as 80 km (50 mi) from water greatly augments its overall range in otherwise marginal areas). (Kingdon 1997, Nowak 1999)

Because of its enormous size, the African elephant can damage its habitat by pushing over trees to obtain edible upper parts of the trees, sometimes modifying the habitat over large areas. In the past, when the species could roam more freely, these environmental modifications presented few problems and in fact may have been necessary for the maintenance of ecosystems that could support large and diversified populations of animals. As human populations increased in Africa, however, and as large areas were cleared for agricultural purposes, the range of the elephant became fragmented and more confined to restricted sites such as parks and reserves. Unable to migrate or disperse naturally, a growing population of elephants can begin to damage its habitat. (Nowak 1999)  

The African elephant is found in both the Cape Floristic Region, Coastal Forests of Eastern Africa, Eastern Afromontane, Guinean Forests of West Africa, Horn of Africa, Maputaland-Pondoland-Albany, and Succulent Karroo Biodiversity Hotspots (Cons. Intl. 2005) as well as the East African Acacia Savannas, Kaokoveld Desert, Zambezian Flooded Savannas, Sudanian Savannas, and Western Congo Basin Forests Global 200 Ecoregions. (Olson & Dinerstein 1998, Olson & Dinerstein 1999)

Age to Maturity:

The age at sexual maturity in elephants is highly variable and is dependent on habitat conditions and elephant population density. Studies of a number of elephant populations have shown that the population average of the age at sexual maturity ranges from 12 - 23 years. This is the widest range of variability in this characteristic that has been reported for wild mammals. The earliest reported age of female sexual maturity for an individual elephant is 7 years. Under optimal conditions, females usually attain sexual maturity at 11 years. In areas with relatively harsh conditions, females may not become sexually mature until 18 or 19, up to 22 years. (Laws 1981. Moss 1988, Nowak 1999, Whitehouse & Hall-Martin 2000)

Under optimal conditions, males attain sexual maturity when they are about 10 years old; however, although a teenage male is sexually mature, he has to go through a long period of growth and social development before he is able to mate with females. From about 25 years of age, males may come into a physiological state called ‘musth’, an Urdu word meaning ‘intoxicated’. It is the phase of heightened sexual activity and aggression in a male elephant’s sexual cycle. Males begin to reproduce when they are 25 - 30 years old and only reach prime breeding age between 40 - 50 years old. (Poole & Thomsen 1989, Moss 1992, Nowak 1999)

Examples of the Age to Maturity:

  • Budongo Forest Reserve (Uganda): The age of puberty for females was delayed until they were 22 years old (Moss 1988).
  • Addo Elephant National Park (South Africa): The earliest age at which females have produced their first calf is 10 years; by 16 years of age, 95 % of cows have had at least one calf. The mean age of first calving is 13.0 years (SD: 2.0 years (n = 66)). (Whitehouse & Hall-Martin 2000)
  • Amboseli (Kenya): 11 - 12 years is the common age for puberty in females, although there were a few who started cycling earlier, at 9 and 10 years. At the same time there were some females who did not appear to give birth for the first time until they were over 20 years old. Male calves left the family on reaching sexual maturity at anywhere from 10 - 15 years of age. (Moss 1988)
  • Mkomazi Game Reserve (Tanzania): Females were reaching puberty at the age of 12 years (Moss 1988).
  • Lake Manyara (Tanzania): Females reached puberty at 11 (Moss 1988).

Gestation Period:

The average gestation period of the African elephant is 22 months, with a recorded range of 17 - 25 months (Nowak 1999).  This is the longest pregnancy among living mammals (Ricciuti 1993).

Birth Season:

Births may occur at any time of year, but there is a calving peak just before the height of the rainy season in the case of the savanna elephant. This provides the young elephants with a cool environment and an abundance of good cover. Savanna cows cease to ovulate during the dry season when food is not as abundant, nor of such high nutritional value, thus ensuring that the calf is born in the wet season. In the case of forest elephants, where food abundance and quality are less variable, breeding seasons are not as clearly defined. (Stuart & Stuart 1996, Nowak 1999)

Birth Rate and Population Growth:

Females produce calves at intervals of 2.5 - 9 years (usually about 5 years). There is normally a single young, but twins occur in 1 - 2 % of births (Nowak 1999).

Population Growth Rate: An African elephant population can increase at an annual rate of 4 - 5% under favorable conditions, the maximum being 7 %.  A maximum growth rate of 6 - 7 % can be achieved if, for example, no elephant dies until it reaches old age, females start conceiving at 9 or 10 years of age, and they give birth to a calf every 3 years. (Moss 1992. Nowak 1999)

In a study in Addo Elephant National Park, South Africa, of the total number of sexually mature cows under the age of 49 (n = 75), 95 % were observed to be breeding (pregnant or lactating). (Whitehouse & Hall-Martin 2000

The peak in fecundity among female elephants has been found to occur at different ages in different populations. For example, this peak occurred in 18 - 19 year old females in the Luangwa Valley, Zambia; 25 - 29 year old females in Addo Elephant National Park, South Africa; and 31 - 35 year old females in Northern Bunyoro, Uganda. (Whitehouse & Hall-Martin 2000

Females may average four calves during their lifetime (range 1 - 9), the number dependent on habitat conditions and elephant density (Laursen & Bekoff 1978).

The generation time for African elephants, calculated as the average age of reproductive females, is 25 years (IUCN 2004).

Early Development:

Birth takes place away from the family. The mother, and frequently another female, go off alone and prepare a soft spot in the ground by loosening the dirt with their forefeet. The mother stands during birth and drops the calf head first, rupturing the umbilical cord in the process. The newborn calf weighs 90 - 120 kg (200 - 260 lb). Within 15 - 30 minutes the newborn can stand and will follow the mother back to the family where it is thoroughly inspected by the other members. (Laursen & Bekoff 1978, Nowak 1999) 

It is very important that a young elephant grow up in the presence of a normal social structure. This includes the tending of infants by individuals that are not the infant’s own mother. In elephant society, these caregivers are immature females ranging in age from about 2 - 11 years, and they play an important role in the rearing of calves. They stand over the calves when they are sleeping, go and get them if they wander away, rescue them from getting caught in a bush or marooned in a swamp, and come to their aid at the slightest cry. This behavior allows the mother to feed and rest in peace for much of the time. It also prepares the young female for the rearing of her own young and is beneficial to the calves. Studies in Amboseli (Kenya) have shown that families with many caregivers have a high calf survival rate. (Moss 1992)

Weaning usually takes place after 6 - 18 months, and calves can survive on solid food before they are 2 years old. Occasionally, calves nurse up to 9 years or until the next calf is born. Calves becomes independent at varying ages. Some scarcely leave their mother’s side until they are nearly 10 years old (an age at which some will already have calves of their own). (Laursen & Bekoff 1978, Moss 1988, Kingdon 1997, Nowak 1999) 

Growth of a young African elephant is fairly rapid up to the time the animal reaches maturity. Growth continues at a slower rate throughout life. (Stuart & Stuart 1996) 

Dispersal:

Males start to exhibit signs of independence at around 8 - 9 years of age, when they might spend a day or two away from the family. The age of independence can be defined as the age when a male spends less than 20 % of his time with his natal family. Using this definition, the average age of independence in Amboseli (Kenya) is 14 years, with some leaving as early as 9 years old and others as late as 18 years old. (Moss 1992)  

Females do not disperse from their family, although as the family grows a subgroup of young adult cows may separate to form a new family. (Stuart & Stuart 1996).

Maximum Reproductive Age:

Female African elephants remain fertile until they are 55 - 60 years old (Nowak 1999). Bull elephants at Amboseli (Kenya) remained reproductively active into their 50's (Moss 1992).

Maximum Age:

African elephants can live to be 60 years old in the wild (more than 80 years old in captivity) (Macdonald 2001).

Apart from premature death caused by humans, disease, predation or injury, the life span of an elephant is determined by the length of time its grinding teeth last.  Once the last teeth in the mouth have worn away, the elephant is unable to chew its food, and it will die [see "The Teeth" below]. (Stuart & Stuart 1996)

Diet:

The African elephant feeds on a wide variety of plants, ranging from grass to the leaves, twigs and flowers of trees. Herbaceous plants are rich in protein and starches and are the favored food of the elephant, when available. The forest elephant also feeds on a wide variety of tree fruits. The elephant uses its tusks to strip the bark off trees, sometimes pushing over a whole tree to gain access to the edible upper parts of the tree. In the wet season, grass forms an important part of the diet, and elephants will travel considerable distances to areas where rain showers have stimulated new growth of grass. The African elephant will wade into swamps, lakes and river shallows to feed on reeds and other water plants. In high montane areas the elephant feeds on tussock grasses and heath. It will also dig for underground roots and rhizomes. The African elephant often needs to supplement its diet with extra salt, either by utilizing salt licks where available or by digging up and exposing salt deposits. (Laursen & Bekoff 1978, Botkin et al. 1981, Macdonald 2001, Stuart & Stuart 1996)

The African elephant consumes about 5% of its body weight in 24 hours. Its daily food intake averages 150 kg (330 lb) and ranges up to 300 kg (660 lb). The quality of food has a significant impact on the quantity consumed: the more nutritious the food, the less is required. Feeding is a constant process with a peak in the morning. (Stuart & Stuart 1996, Kingdon 1997)

Elephants generally drink once a day, although in some seasons and in some habitats they can adapt to drinking every other day or even every third day.  During dry periods the elephant excavates holes in dry river beds with its tusks, trunk and feet to gain access to underground water. The daily water intake is 100 - 220 liters (106 - 223 qt).  Forest elephants obtain a significant amount of water from the fruits they eat. In the dry season, the water provided by fruits may be insufficient and these elephants come out of the forest in search of water.  (Moss 1992, Stuart & Stuart 1996, Parren et al. 2002)

Physical and Behavioral Characteristics:

The Trunk: One of the African elephant's most distinctive features is the trunk, a flexible elongation of the upper lip and nose that consists of thousands of muscles. The African elephant has two opposing, fingerlike outgrowths (called "processes") at the tip of its trunk. With these processes and the trunk’s flexibility, the elephant has enough maneuverability to pick up small objects such as berries. The trunk is used for breathing (along with the mouth) and to pick grass, pull down branches, and lift logs; to suck up and squirt water into its mouth for drinking or over its body for cooling; and as a snorkel when the animal is crossing a deep river.

The Tusks: In both sexes one incisor tooth on each side of the upper jaw is greatly developed to form a tusk. The tusks are large, thick, and curved in most bull elephants; cows have smaller tusks. The tusks continue to grow throughout the elephant's life. They are used for stripping bark from trees, digging for roots, excavating for minerals (including salt), and defense. (Stuart & Stuart 1996, Nowak 1999) 

The Teeth: During its lifetime, the African elephant has a succession of a total of six grinding teeth on each side of its upper and lower jaws. This is the only kind of tooth the elephant has (not counting the tusks). These teeth are very large (weighing 3.7 kg (8.1 lb)). The elephant’s teeth do not replace one another vertically, as is usually the case with mammals, but they come in successively from the back of the mouth, the group of teeth gradually moving forward. As the elephant grinds its food, its teeth are worn down. The oldest teeth, in front, become so worn down as to be of no further use. They eventually are pushed out and replaced by the teeth behind them. After the last of these teeth are worn down, the elephant will die of starvation. (Stuart & Stuart 1996, Nowak 1999) 

The Ears: The African elephant’s large, fan-shaped ears are use for cooling.  The undersides of the ears have an extensive supply of blood vessels. Ear flapping, characteristically seen when elephants stand in the shade on hot days, creates air currents over the blood vessels and promotes the loss of excess body heat.  In aggressive displays, the ears are spread out to the side.

Communication by Sound: African elephants communicate with each other by means of visual signals and touch, but they are also very vocal animals. They have an extensive and varied repertoire of vocalizations, ranging from soft rumbles to ear-splitting trumpets. The most common calls are the various rumbles, and research has shown that all of these have infrasonic components. This explains a surprising phenomenon that has long been observed: African elephants are able to communicate when not within sight of each other, at distances over which sounds audible to humans would die out. For example, a herd which is feeding quietly may suddenly become alert and take flight for no apparent reason. These low frequency sounds (14 - 35 Hz) can carry over relatively long distances, possibly as far as 10 km (6 miles). (Moss 1988, Moss 1992, Stuart & Stuart 1996) 

Activity Patterns: Activity is both diurnal and nocturnal but drops during the hottest hours of the day. One study reported the following estimates of the percentage of time spent by African elephants in different activities over 24 hours: resting - 13%, feeding - 74%, traveling - 11%, other - 2% (Schaller et al. 1985).

Bathing: Despite the thickness of an African elephant's skin (up to 3 cm (1.2") thick), its skin is very sensitive and requires constant care. To keep its skin healthy, the African elephant takes baths in water, wallows in mud, and takes dust baths. It  sucks dust up into its trunk and blows it out through the trunk, thus depositing the dust on its back and head. Dust and mud act as sunscreens, protecting the elephant’s skin from the sun. They are also good insect repellents, deterring insects from biting the sensitive skin. (Laursen & Bekoff 1978, Ricciuti 1993, Burnie & Wilson 2001) 

Sleeping: An eyewitness account: "The elephants fed continuously until midday, when they moved to higher, dry ground, where they found a bare patch of dusty soil... Now they gathered together in a tight group, nearly a circle, standing close together, some touching... First one calf lay down, then three more subsided to the ground and lay flat on their sides. The females’ heads hung down and their trunks became limp and stretched out until the tips touched the ground. Two of the females rested their trunks on their tusks... A half-grown calf placed his trunk on the back of a sleeping calf. All became quiet and breathing deepened. The elephants slept for about 40 minutes..." (Moss 1988)

Movement: The African elephant walks at speeds of 2 - 6 kph (1 - 4 mph) and can speed up to a shuffling gait of 16 - 24 kph (10 - 15 mph). When charging the elephant can move at a rate of 35 - 40 kph (22 - 25 mph) over a short distance. Daily average movements of 12 km (7 mi) have been reported, and when searching for food and water, the African elephant can walk up to 500 km (300 mi). Although it is sure-footed, because of its large size and weight the African elephant moves carefully and will generally not attempt to step across a ditch more than 1.5 m (5') wide. When it encounters open water, an elephant can submerge with only the nostrils at the tip of its trunk showing above the surface. In this manner an elephant can swim or walk underwater. (Laursen & Bekoff 1978, Ricciuti 1993, Stuart & Stuart 1996, Nowak 1999) 

Migration: Elephants do not travel far if food, water, and shade all remain available. If these items become scarce, elephants will travel long distances to find an adequate supply (as long as their movements are not restricted because of man's activities). In the past, African elephants made annual migrations of up to 300 km (200 mi) to find desirable habitat. Nearly all seasonal movements followed the same general pattern: a migration from permanent water sources at the start of the rainy season, followed by a movement back to permanent water when the rains and temporary water holes dried up. Often there was a concentration of animals in suitable areas during the dry season or in times of drought and a dispersal in the wet season. Shorter, nonseasonal movements of elephants between water and feeding areas occurred. The animals followed regular routes and sometimes created well-worn paths. At present, the African elephant is often forced into unnatural concentrations in protected areas, and migration is restricted. (Laursen & Bekoff 1978, Nowak 1999) 

Reproductive Flexibility: Studies of elephant populations at different population densities and habitat conditions have shown that the elephant populations appear to have a built-in birth control system. There are significant differences in reproductive parameters between elephant populations, and these differences can lead to profound differences in population growth rates.  For example, the age of sexual maturity for females can be delayed and the average interval between the births of calves can be lengthened under an adverse relationship between habitat conditions (e.g. low food supply and/or high grazer competition) and a relatively high elephant population density, to the extent that the population will decrease over time. At the opposite extreme, during times of plentiful food supply with low population density and competition, the population could reduce the age of sexual maturity for females and shorten the calving interval to achieve a higher growth rate (See Age to Maturity and Birth Rate and Population Growth above.). (Croze et al. 1981, Moss 1988, Moss 1992) 

Elephant Graveyards:  "There is a myth that elephants that are ill go off to an ‘elephant graveyard’ to die. Although the myth is not true, it is also not without some foundation. Probably in any given elephant range there are places that sick and wounded elephants tend to go. These would be areas where there is water and shade and soft vegetation to eat. Places such as this might have more carcasses than other parts of a population’s range, and therefore people may have thought there was a special area where elephants went to die. More likely the myth arose from hunting practices that involved killing several elephants at the same time... [such as] by building a ring of fire around them. In this case the bones of many elephants would have been found in one place." (Moss 1988) 

Social Organization:

The Family: The major unit of elephant society is a stable, matriarchal family of cows and their calves. A typical family in an unpoached population includes an average of 10 individuals, usually consisting of an older and experienced female (the matriarch), her dependent offspring, and her adult daughters with their immature offspring. In forests, such families may include only one or two offspring. The matriarch usually leads the family until death, when she is succeeded by her eldest daughter. Occasionally the family may include one of the matriarch’s sisters and her offspring as well. Members of a family work together to take care of the calves and to signal each other of impending danger. Males are usually driven out of the family when they attain puberty (usually at 10 - 15 years of age), after which they remain solitary or temporarily congregate into bachelor herds where no long-term social bonds exist between the different individuals. They only associate with families when one of the females is in estrus. Sometimes families simply get too large to be efficient social groupings, and either they split more or less down the middle or a subgroup breaks off. (Laursen & Bekoff 1978, Moss 1988, Kingdon 1997, Nowak 1999, Nyakaana et al. 2001) 

Relations between Family Members: Communication between family members takes many forms, including vocalizations, body postures, and frequent touching of each other with their trunks. There is a dominance hierarchy based on age, but more obvious is the cooperative behavior among elephants, such as the care of calves by members of the family other than their mothers, particularly adolescent females. Cooperative behavior also includes providing mutual defense against predators and employing a system of lookouts while bathing. Elephants have even been reported to assist wounded comrades by supporting them between two adults and helping them to move away from danger.  (Laursen & Bekoff 1978, Moss 1988, Moss 1992, Burnie & Wilson 2001) 

Larger Groups: Aerial counts of elephant families at Lake Manyara (Tanzania) showed that, far from avoiding each other, the families were always more clumped than would be expected by chance movements. This clumping can be seen in other areas such as the Serengeti (Tanzania), where one can fly for hours without seeing an elephant, then suddenly encounter 10 - 15 families within the area of a square mile. In fact, African elephants appear to have a social affinity for one another and can be found in larger groups of various sizes. The groups may include hundreds of individuals and are usually temporary in nature, although some families appear to have special relationships with certain other families in a population and spend a great deal of their time together. The aggregations have mostly been observed among savanna elephants in East Africa. It is rare to see more than 10 forest elephants together, although groups of over 100 forest elephants have been observed at certain salt licks in the Central African Republic. (Douglas-Hamilton & Douglas-Hamilton 1975, Moss 1988, White 1993, Nowak 1999) 

The Greeting Ceremony: The best indicator of the strength of the bonds between elephants is the greeting ceremony. Almost all elephants will greet one another, but the nature and intensity of the greeting depends on who the elephants are, what their relationship is, and if it is a close one, how long they have been separated. A mild greeting between several elephants may involve raising and flapping their ears, rumbling the throaty "greeting rumble," and "exchanging trunks"; i.e., putting the tips of their trunks in each other’s mouths. If two subgroups of a family have been separated for a few days, the greeting ceremony will be far more intense. They will run together, rumbling, trumpeting, and screaming; click their tusks together; entwine their trunks; flap their ears; spin around and back into each other; urinate and defecate; and generally show great excitement. A greeting such as this will sometimes last for as long as 10 minutes. (Moss 1988, Moss 1992) 

Aggression: Aggression is shown by raising the head and trunk, extending the ears perpendicular to the body, kicking dust, swaying the head, and making either a mock or a serious charge. Most dominance struggles are resolved after some pushing and light tusking, but battles between males for mating privileges sometimes involve fatal use of the tusks. (Nowak 1999)  

Gender Distribution:

Studies in Addo Elephant National Park (South Africa) from 1954 - 1998 showed that the overall sex ratio of newborn calves did not differ significantly from 1:1 (1954 - 1998 births: 157 males, 163 females) (Whitehouse & Kerley 2002). Studies in Amboseli (Kenya) between 1978 - 1980 also showed a sex ratio very close to 1:1 among newborns, with a total of 50 males and 49 females being born (Moss 1988).

Mortality and Survival:

General Findings

  • Man is the only predator that causes mortality that has a limiting effect on elephant populations. Other predators, including lions, wild dogs, hyenas and crocodiles, usually prey only on isolated calves less than 2 years old. (Laursen & Bekoff 1978)  
  • About half of all elephants die before they reach the age of 15 (Stuart & Stuart 1996)
  • Between 2.5 - 5% of the females of all ages in an elephant population die each year (Whyte et al. 1998)
  • Cumulative mortality during the first 3 years is about 27 - 38%, dropping to about 3 - 3.5 % annually for the ages from 3 - 45 years and rising again to about 10 - 20 % after that. (Laursen & Bekoff 1978, Wu & Botkin 1980) 

An Example: Addo Elephant National Park (South Africa), 1976 - 1998: Mortality rates for calves in their first year are similar for the two sexes, but thereafter male mortality rates are higher than those of females in all age classes. This results in a higher percentage of females than males in the overall population. (Details: Average annual mortality rates in the following age classes (M=male, F=female): age class 0: M: 6.2 %, F: 6.2 %; age class 1-9: M: 0.9 %, F: 0.1 %; age class 10-19: M: 2.0 %, F: 0.4 %; age class 20-29: M: 3.1 %, F: 0.3 %; age class 30-44: M: 5.1 %, F: 1.2 %; age class 45-59: M: 100 %, F: 1.6 %; age class 60-63: F: 100 %.) Following the fencing of the park in 1954, the majority of adult cows died of natural causes. The primary cause of death among adult bulls, however, was intraspecific fighting, and the death rate from this fighting is unusually high compared to other elephant populations. That suggests that confinement of elephants in the park by fencing may have had a negative impact on the social behavior and mortality of adult males. (Whitehouse & Kerley 2002) 

Density and Home Range:

Examples of African elephant density:

  • Cameroon: Lobeke Forest Reserve: The estimated density of forest elephants in the Lobeke forest reserve, 4 - 6 elephants/sq km (10 - 16 elephants/sq mi), is reportedly the highest in the Central African forest region (Usongo 1998).
  • Gabon: Lowland forests: Elephant densities in the lowland forests of Gabon were estimated to be 0.4 elephants/sq km (1 elephant/sq mi) in undisturbed lowland forest and 0.1elephants/sq km (0.3 elephants/sq mi) in disturbed lowland forest (Cumming et al. 1990).
  • Kenya: Amboseli National Park: A relatively undisturbed population of about 500 elephants had an overall density of 0.14 elephants/sq km (0.36 elephants/sq mi), increasing during the dry season to 0.4 - 0.9 elephants/sq km (1.0 - 2.3 elephants/sq mi) (Nowak 1999).
  • South Africa: Addo Elephant National Park: Between 1954 and 1998, population density was 1.0 - 4.0 elephants/sq km (2.7 - 10.4 elephants/sq mi) in Addo, which was totally enclosed by an elephant-proof fence (Whitehouse & Kerley 2002).
  • Tanzania: Lake Manyara: About 5.4 elephants/sq km (14 elephants/sq mi) (Moss 1988).
  • Uganda: Murchison Falls area: 0.8 - 1.2 elephants/sq km (2 - 3 elephants/sq mi) (Fowler & Smith 1973).

Home range:

The size of the home range depends on the abundance and nutritional value of food, the availability of water and, in the case of savanna-dwelling animals, shade. Home ranges in arid areas are considerably larger than those in well-watered areas. Where conditions are ideal, the home ranges of families may be from 15 sq km (5.8 sq mi) to 50 sq km (19 sq mi). The home ranges of bull elephants are considerably larger (up to 1500 sq km (580 sq mi)), since bulls move around seeking cows that are receptive to mating. Home ranges of the forest elephant are usually considerably smaller than those of the savanna elephant, mainly because of the abundance of food and the ready availability of water in the forest habitat as compared to the savanna habitat. (Stuart & Stuart 1996) 

A study in Lake Manyara National Park (Tanzania) found that the home ranges of elephant families overlapped extensively with one another. There was no indication of territoriality, but each family did not roam equally throughout the whole park. Most of them preferred certain areas. Most families traveled through most of their home range every month. (Douglas-Hamilton & Douglas-Hamilton 1975) 

Detailed studies of home ranges of the forest elephant were initiated around the beginning of the 21st century, and first results indicate that the home ranges can be as much as 60 km (37 mi) across, far larger than previously thought (Macdonald 2001).

The Kaokoveld, an area of the Namib Desert in Namibia, is rugged and mountainous, with open gravel plains bisected by dry riverbeds. It contains elephants at the edge of their natural range, with the largest home ranges recorded anywhere in Africa: 5800 - 8700 sq km (2200 - 3400 sq mi). (Berger 1997) 

The following list provides published home range sizes of male and female elephants based on minimum convex polygons, as well as annual rainfall for each location (home ranges are for female-led families unless they are indicated to be for males) (NP = National Park) (R = rainfall) (Osborn 2004):

  • Amboseli NP (Kenya): 2800 sq km (1080 sq mi); R = 350 mm (14")
  • Hwange NP (Botswana): 1040 - 2500 sq km (400 - 970 sq mi); R = 630 mm (25")
  • Hwange NP (Botswana) (male): 1300 - 3000 sq km (500 - 1200 sq mi); R = 630 mm (25")
  • Kruger NP (South Africa): 130 - 1260 sq km (50 - 490 sq mi); R = 550 mm (22")
  • Laikipia (Kenya): 450 - 500 sq km (170 - 190 sq mi); R = 750 mm (30")
  • Laikipia (Kenya): 600 - 800 sq km (230 - 310 sq mi); R = 400 mm (16")
  • Lake Manyara NP (Tanzania): 10 - 57 sq km (3.9 - 22 sq mi); R = 1000 mm (39")
  • Namibia: 5800 - 8700 sq km (2200 - 3400 sq mi); R = 320 mm (13")
  • Queen Elizabeth NP (Uganda): 360 sq km (140 sq mi); R = 900 mm (35")
  • Queen Elizabeth NP (Uganda) (male): 500 sq km (190 sq mi); R = 900 mm (35")
  • Sengwa (Zimbabwe) (male): 320 sq km (120 sq mi); R = 670 mm (26")
  • Transvaal (South Africa): 120 - 470 sq km (46 - 180 sq mi); R = 600 mm (24")
  • Transvaal (South Africa) (male): 160 - 340 sq km (62 - 130 sq mi); R = 600 mm (24")
  • Tsavo East NP (Kenya): 2400 sq km (930 sq mi); R = 300 mm (12")
  • Tsavo East NP (Kenya) (male): 1040 - 1210 sq km (400 - 470 sq mi); R = 300 mm (12")
  • Tsavo West NP (Kenya): 410 sq km (160 sq mi); R = 550 mm (22")
  • Tsavo West NP (Kenya) (male): 290 - 340 sq km (110 sq mi); R = 550 mm (22")
  • Waza NP (Cameroon): 2500 - 3100 sq km (970 - 1200 sq mi); R = 700 mm (28")
  • Zambezi Valley (Zimbabwe): 160 sq km (60 sq mi); R = 800 mm (32")

References

Barnes et al. 1999, Berger 1997, Blanc et al. 2003, Botkin et al. 1981, Brookfield Zoo, Burnie & Wilson 2001, Chadwick 1991, CITES, Cons. Intl. 2005, Croze et al. 1981, Cumming et al. 1990, Czech Web Site, Douglas-Hamilton & Douglas-Hamilton 1975, Eggert et al. 2002, Fowler & Smith 1973, Groning and Saller 1998, Hardman 2002, IUCN 2000, IUCN 2004, Kingdon 1997, La Monte & Welch 1934, Laursen & Bekoff 1978, Laws 1981, Leader-Williams 1996, Macdonald 2001, Moss 1988, Moss 1992, Naturefoto-Online, Nature-Wildlife, Nowak 1999, Nyakaana et al. 2001, Olson & Dinerstein 1998, Olson & Dinerstein 1999, Oryx 1978f, Oryx 1980d, Oryx 1981d, Osborn 2004, Parker & Martin 1982, Parren et al. 2002, Poole & Thomsen 1989, Ricciuti 1993, Said et al. 1995, Schaller et al. 1985, Sharp 1997, Sharp 1997b, Stuart & Stuart 1996, Usongo 1998, White 1993, Whitehouse & Hall-Martin 2000, Whitehouse & Kerley 2002, Whyte et al. 1998, Wood 1860, Wu & Botkin 1980, WWF, WWF 1997


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