African elephant

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African elephant
African elephant (Loxodonta africana)

African elephant ( Loxodonta africana )

Systematics
without rank: Paenungulata
without rank: Tethytheria
Order : Russell animals (Proboscidea)
Family : Elephants (Elephantidae)
Genre : African elephants ( Loxodonta )
Type : African elephant
Scientific name
Loxodonta africana
( Blumenbach , 1797)

The African elephant ( Loxodonta africana ), also African steppe elephant or African bush elephant , is a species from the elephant family . It is the largest land mammal currently living and at the same time the largest recent land animal on earth. In addition to the tusks and the distinctive trunk, the large ears and columnar legs are outstanding features . The African elephant differs from its somewhat smaller relative, the elephant, in numerous morphological and anatomical featuresForest elephants and the Asian elephant . The distribution now includes large parts of Africa south of the Sahara . The animals have adapted to numerous different habitats there, ranging from closed forests and open savannah landscapes to swamp areas and desert-like regions. Overall, however, the occurrence is highly fragmented.

The way of life of the African elephant has been well researched through intensive studies. It is characterized by a strong social character. Female animals and their offspring live in family groups (herds). These in turn form a more closely related clan. The individual herds meet on certain occasions and then separate again. The males form bachelor groups. The various associations use action spaces in which they sometimes wander around in an annual cycle. The animals use different tones in the low frequency range to communicate with one another . The individual individuals can recognize one another based on the sound produced, but also through certain chemical signals. In addition, there is an extensive repertoire of gestures. The cognitive abilities of the African elephant should also be emphasized .

The diet consists of both soft and hard vegetable foods. The composition varies regionally and seasonally. In general, the African elephant spends a large part of its daily activities eating. Reproduction takes place all year round, regionally there are tendencies towards stronger seasonalization. Bulls come to the Musth once a year , during which they go on a hike in search of cows willing to breed. During the musth the aggressiveness is increased, rival fights take place. The cows' sexual cycle is comparatively long and atypical for mammals. After the birth, he usually takes several years off. Usually, after a gestation period of almost two years, a young animal is born that grows up in the maternal herd. Young females later remain in the herd, the young males leave it.

The first scientific description of the African elephant took place in 1797 with a formal separation of the African from the Asian elephant. The generic name in use today, Loxodonta , was not officially introduced until thirty years later. The name refers to the distinctive tooth differences between the Asian and African elephants. During the 20th century several subspecies were distinguished, including the forest elephant of central Africa. According to genetic studies, the latter is now considered an independent species; the other subspecies are not recognized. In tribal history, the African elephant can be documented for the first time in the beginning of the Middle Pleistocene . The entire population is considered to be endangered. The main reasons for this are the hunt for ivory and habitat loss due to the increasing human population. The African elephant is one of the so-called " Big Five " of big game hunting and safari.

features

Habitus

The African elephant is the largest living land mammal and is relatively easy to recognize by its trunk , tusks , large ears and columnar legs. The head-trunk length is 600 to 750 cm, the tail is 100 to 150 cm long. According to studies in the Amboseli National Park, an adult bull has a shoulder height of 290 to 370 cm. Cows are on average smaller, measuring between 250 and 300 cm. For the Kruger National Park , the size values ​​for males are a maximum of 345 cm, for females 274 cm, whereby 255 cm is rarely exceeded. Maximum weights for bulls are given as 6048 kg, for cows with 3232 kg. One particularly large individual weighed 6,569 kg. The largest scientifically measured specimen, an animal from Fenykoevi in Angola , had a shoulder height of 400 cm and a weight of around 10 tons, it is on display today in the Smithsonian Institution in Washington, DC .

The head is massive and large, broader in adult bulls than in cows and younger individuals. In contrast to the domed, two-humped forehead of the Asian elephant ( Elephas maximus ), the African elephant has a flat, receding forehead. The characteristic large ears measure around 120 cm in width and up to 200 cm in height. Its outline is somewhat reminiscent of the contours of the African continent. They are significantly larger compared to the Asian elephant. The trunk ends in two " fingers ", the Asian elephant has only one. Another difference to the Asian elephant can be found in the back course, which is more saddled in the African elephant, so that the highest point is reached on the shoulders. The Asian elephant has a characteristic hump back, the highest point on the body is also on the forehead. The forest elephant ( Loxodonta cyclotis ), on the other hand, has a straight back line. The general skin color corresponds to a pale gray to brown tone, in rare cases light, pigment-free spots can be seen. The thickness of the skin reaches up to 40 mm on the legs, the front head and the back, and is generally wrinkled. The body hair is sparsely distributed over the body. Young animals, however, often have a loose, reddish-brown fur. In adult animals, longer hairs are usually only formed on the chin and trunk; they are around 40 mm long. In addition, there is a tuft of hair up to 50 cm long, sometimes blackish, shiny hair at the end of the tail. The front and rear feet each have five toes. Externally there are four to five hoof-like nails at the front and three to five at the back. The sole is soft and cracked and individually designed. It leaves characteristic rounded traces. Those of the front feet generally exceed those of the hind feet. The length of the hind feet varies from 34 to 54 cm, it is proportional to the shoulder height of an animal.

Skull and dentition features

African elephant skull; The nostril in the middle and the alveoli of the tusks below are clearly visible.

The skull is generally built large, a measured specimen had a length of 92 cm and a width of 73 cm at the zygomatic arches . Compared to the Asian elephant, the skull of the African elephant is significantly more rounded and less high. When viewed from the front, the forehead line of the African animals is straight or arched and not as markedly indented as that of the Asian. The extensive air-filled chambers in the skullcap, which are reminiscent of honeycombs, are typical of elephant skulls . On the one hand, they reduce the weight of the skull, but on the other hand they also increase the surface area, which means that more attachment surfaces are available for the massive neck and masticatory muscles. Due to the extreme pneumatization, the skullcap can be up to 30 or 40 cm thick. The facial area of ​​the skull is made up of the nasal bones , the upper jaw , the zygomatic bone and the median jawbone . The nasal bone is small due to the trunk formation, the nostril is at the level of the orbit and thus lower than in the Asian elephant. The middle jawbone forms the outer shell of the alveoli of the tusks. In the African elephant the alveoli are laterally apart and protrude slightly curved forwards, in the Asian elephant, on the other hand, they are directed downwards. As a general characteristic of Tethytheria , the middle jawbone fuses with the frontal bone, among other things . The frontal bone itself is very broad and rounded in the African elephant, but narrow in the Asian elephant because it moves in above the nostril. The parietal bones represent the largest skull bones. A small furrow runs down on them, which is more pronounced in the Asian elephant. On the occiput , the articular surfaces for articulation with the cervical spine clearly point backwards. They are also at the level of the orbit and thus lower than their Asian counterpart. The external auditory canal is located in the immediate vicinity of the occipital joints, while it is positioned lower in the Asian elephant.

The lower jaw has a rather delicate structure with a long and low horizontal bone body. It differs from the broad and swollen lower jaw of the Asian elephant. When viewed from above, it is V-shaped, while that of the Asian elephant looks more U-shaped. Overall, the lower jaw consists of massive, compact bones. The symphysis on the anterior lower jaw, connecting the two mandibular arches, extends forward in a long and narrow process. With an average length of 17 cm, the bone connection itself is relatively and absolutely shorter than in the smaller forest elephant. Several openings are formed on the outside, which are referred to as the lateral and central foramen mental . The rear end of the lower jaw appears rounded due to the curved angular process. The ascending branch is wide, the crown process rises only a little above the occlusal plane and lies about half the length of the entire lower jaw. It is straight in shape in contrast to the rounded crown of the Asian elephant. The articular process, on the other hand, runs straight up and not angled inward as in its Asian relative. A large mandibular foramen opens on the inside of the ascending branch . The outside, on the other hand, hides the flat indented masseteric fossa , which is not as deep as that of the Asian elephant.

The dentition consists of 26 teeth together, committed to following dental formula group: . The tusks , which are only formed in the upper dentition and consist of the hypertrophied second incisors, are characteristic. In the African elephant, in contrast to the Asian elephant, they usually occur in both male and female animals. The tusks have a clearly curved and partly outwardly curved course, different from those of the forest elephant with their rather straight shape. Record measurements reach up to 345 cm. An extremely large pair of tusks comes from an individual from Kilimanjaro with lengths of 319 or 311 cm and a weight of 97 or 102 kg. In the Kruger National Park, the heaviest tusk of a cow to date weighed around 11 kg, but weights over 7 kg are rarely reached. In old bulls, the greatest weight registered so far was 73.5 and 69 kg, respectively. The basal diameter of the tusks of male animals varies between 15.5 to 19.6 cm, that of female animals between 8 and 11.9 cm. In the bull, they grow both in length and in width for a lifetime. In contrast, the tusks of the cow after the age of 15 only develop in length and this at a much slower rate than in the bull; they therefore appear slimmer than those of the males. Often there is a size discrepancy between the left and right tusk of an individual, which is due to more intensive use of one of the two tusks. This different usage refers to a certain "tuskedness" (in the sense of handedness ) in the African elephant. The tusks are already created as “tusks” in the embryoal stage , but do not come through. Later the permanent tusks replace the "tusks".

Comparison of the molars of elephants. Above: African elephant. Middle: Asian elephant. Below: woolly mammoth.

As with all elephants today, the rear dentition consists of three premolars and three molars per half of the jaw. The change of teeth takes place horizontally in that a new tooth is pushed out from behind when the previous one has largely been chewed off. As a result, the African elephant (like the Asian elephant) changes its teeth a total of five times after the eruption of the first tooth (premolar dP2). As a rule, the first tooth is already functional at birth and falls out at 1 to 2 years. The eruption of the other teeth occurs for the premolars at 1.5 (premolar dP3) and 2 years (premolar dP4) and for the molars at 5 (molar M1), 15 (molar M2) and 23 years (molar M3). The corresponding loss occurs at 3 to 4 (dP3), 9 to 10 (dP4), 19 to 25 (M1), 43 (M2) and around 65 (M3) years. However, there is a high individual range here. The horizontal change of teeth also has the effect that only one to a maximum of one and a half teeth are in use at the same time per jaw half. In the entire dental structure, the African elephant is the more conservative form compared to the Asian elephant. The teeth generally have lower crowns than their Asian cousins. As with all representatives of the elephant, both the premolars and the molars consist of several lamellar enamel folds . The number varies between the teeth, the second premolar consists of five enamel folds, the third molar of an average of 13 with fluctuations between 14 and 16. The lamella frequency is thus an average of 5.5 (number of lamellae on 10 cm tooth length). Both features are significantly lower than in the Asian elephant. In contrast, the enamel of each bar is markedly thicker with 2 to 5 mm. In the plan view of the Kauoberfläche the enamel each melt bulges fold approximately in the middle rhombic outwardly as it extends substantially parallel in Asiatic elephant. The last molar is the largest molar in the dentition. It can be up to 21 cm long and weigh around 3.7 kg. The first premolar, on the other hand, is only 2.4 cm long and weighs just under 10 g.

distribution

Distribution of the African elephants (genus Loxodonta ) in 2007

The original range of the African elephant encompassed the entire African continent and once also included northern Africa to the Mediterranean coast and southern Africa to the Cape of Good Hope . Various rock paintings and ancient historical reports give evidence of this . Climatic fluctuations in the course of the Holocene displaced the species from the extremely dry landscapes. The northern populations eventually disappeared in the first centuries of today's calendar. Today's occurrence is limited to sub-Saharan Africa . According to estimates, the African elephants now inhabit an area of ​​almost 5.2 million square kilometers, of which slightly more than half is in Central and West Africa , where the forest elephant is also found. As a result, the current range of African elephants only takes up about 20% of the former habitat. The populated landscapes are very fragmented and spotty, individual areas are rarely visited by the African elephant. The decline in the distribution of the African elephant is mainly due to ivory poaching and habitat destruction, including the areas in West and South Africa. In Mali near Gourma there is a small population of around three hundred desert-dwelling elephants, a larger group of over 3000 individuals can be found in the WAP national park complex . Further northern populations can be found in Chad , which also represents the northern limit of today's distribution. In bordering Central Africa and in southern West Africa, the African elephant is being replaced by the smaller forest elephant. Among other things, there is a more or less wide corridor in which the two species hybridize with each other . The African elephant is relatively common in East Africa , where it occurs in numerous fragmented habitats in Ethiopia , Uganda , Rwanda , Kenya and Tanzania . The largest contiguous population is found in the northern part of southern Africa and stretches from Namibia and Angola in the west eastwards across Botswana , Zambia and Zimbabwe . The African elephant has largely disappeared from the southern part of southern Africa. However, he was reintroduced in various places by animals from the Kruger National Park .

“Desert elephants” in the Huab River

The African elephant has adapted to a wide variety of habitats. These consist of semi-deserts , open grass and savannah landscapes , floodplains or swamps as well as a variety of different forest biotopes such as gallery forests , mountain forests or tropical lowland rainforests . The prerequisites for the presence of the African elephant are sufficient water and food. Shade or retreat areas for protection also play an important role in the spread. Elephants do not usually live in deserts . Exceptions are the already mentioned populations of the southern Sahara in Mali and the border areas to the Namib , such as Kaokoveld and Damaraland , in Namibia , where there are also some populations of "desert elephants". In the mountains, the African elephant can occasionally be found at heights of up to 4875 m, such as on Kilimanjaro ; its preferred habitat, however, is in the lowlands.

The strongly fragmented distribution area of ​​the African elephant results in different population sizes and densities. In open grasslands the number of animals is about 0.5 to 2 individuals per square kilometer; it can increase to about 5 per square kilometer in areas with woodland with abundant food supply, such as in Lake Manyara National Park . For ecological reasons, however, such a high population density can be viewed as rather unstable over a longer period of time. Dense forests or deserts, on the other hand, usually only have a small population of animals. Estimates of the total number of individuals in Africa are difficult, since they require a clear separation, at least for those countries with a simultaneous occurrence of the forest elephant and / or hybrids between the two species. For 2007 the numbers were around 496,000 individuals in western, eastern and southern Africa, and for 1991 it was roughly 267,000 to 372,000. The Great Elephant Census produced a total of 352,000 animals in 18 countries in 2014, which is a decrease of 30% compared to 2007. The largest populations are found in the northern part of southern Africa, i.e. the area with the largest contiguous population, where about 234,000 animals live, followed by eastern Africa with around 73,000 animals.

Way of life

Territorial behavior

General and activities

The way of life of the African elephant has been intensively researched, the studies in Amboseli National Park over a period of 35 years should be emphasized. The animals are both diurnal and nocturnal. They spend the majority of 75% of their activity phase eating. As observations in the Ruwenzori National Park in Uganda have shown, there are often three feeding seasons, which are spread over the morning, late afternoon and midnight. The greater part is consumed during the day. The African elephant mostly migrates during the twilight phase. On average, he urinates and defecates about a dozen times a day. There are no major differences in the available time budget between male and female animals. According to studies in the Chobe National Park in Botswana, the average sleep duration is around 2.2 to 2.5 hours within a day, with animals in the Ruwenzori Mountains it takes a little over three hours. As a rule, the animals sleep between 02:00 and 06:00 after midnight. Under certain circumstances, such as malfunctions, you will be awake for two days continuously. This makes the African elephant one of the mammals with the shortest sleep phase. He often stands while sleeping, which means that the REM phase is rarely reached. He only goes into a lying position every three to four days, but this only lasts a little longer than half an hour. As a rule, an animal chooses new sleeping and resting places every day. According to observations, the intensity of daily activity has no influence on the length of sleep.

African elephant skin with clearly visible furrows and crevices in the upper layer; Different skin layers are shown, at the bottom right an example of an Asian elephant

There is also no influence of the activities on the body temperature. The African elephant has a comparatively low body temperature of around 36 ° C compared to large ungulates in the same habitat. The daily variations are relatively small in relation to the sometimes considerably different external conditions. The maximum body temperature in the daily fluctuation rhythm does not come about until relatively late, around 10:00 p.m. In contrast to the Asian elephant , the fluctuating body temperature does not seem to provide a mechanism for thermoregulation . The African elephant inhabits a large number of landscapes, some with severe climatic extremes. To control the heat balance, the animals use, among other things, their ears, which produce both cooling and warming effects by waving and thus represent the largest temperature-regulating organ in the animal world. However, waving one's ears also has an important social function or expresses individual needs and sensitivities. The evaporation of water on the body surface is another option for maintaining body temperature. In contrast to the Asian elephant, the heavily cracked and therefore clearly ornamented skin has a beneficial effect. The water is stored in the furrows and crevices five to ten times longer than directly on the surface. In adult animals, the ornamentation is caused by the cornification and permanent thickening of the skin, which is then torn open by bending stress in the upper epidermis .

Social organization and use of space

Elephant herd with young animals in the Etosha National Park

The social behavior of the African elephant is complex. There are clear differences between male and female individuals. A total of four social units are known in female animals. The base and lowest level (first level) consists of the mother and one or more young animals. Several of these small mother-young animal units form a family group or herd (second level). The family group represents the stable unit in the social community of the African elephant, it is not influenced by seasonal conditions and resources. According to studies in the Samburu National Reserve, it is composed of 2 to 16 individuals. In general it can be said that family groups in more closed landscapes are smaller than in open ones, and the family groups of older cows also outnumber those of younger cows in terms of number of individuals. The members are usually related to each other. Individual herds, including those in areas with heavily human-influenced populations , do not necessarily include animals that are genetically related to one another. At the head of the herd is a mature, mostly older and large lead cow , who not only dominates the other members, but also makes important decisions in critical situations. Your experiences are therefore essential for the family group. The hierarchy within the herd is organized linearly, which minimizes conflicts in the herd and prevents injuries caused by rank fights. Therefore, after the lead cow dies, a high-ranking daughter usually takes over the herd. Larger family groups can then split up into individual daughter communities. With its hierarchically structured herd structure, the African elephant differs significantly from the Asian elephant, in which the hierarchy is much flatter and a lead cow does not play a major role. Several family groups together form a family association or a kinship group (third level). In the Samburu National Reserve, an average of two and a half family groups form a larger family group, it includes 14 to 48 individuals and can span up to four different generations. Here, too, the size and scope depend on the yield of a landscape with smaller kin groups in more wooded regions compared to larger ones in drier areas. The highest organizational unit is the loose clan (fourth level). A clan is characterized by the fact that the individual herds use similar action spaces . The higher levels of social organization above the herd are not stable. The individual herds meet at specific conditions and then separate again, often depending on local conditions and available food sources. In many cases the merger occurs during the rainy season when the food supply increases. The split then takes place during the dry season with dwindling food resources and increasing competition among each other. However, local factors such as the density of potential predators or possibly the social and ecological exchange of information can also influence such associations. Such a social community is known as a fission-fusion social community (“separating and coming together”). The members of the clans are able to differentiate between closely and distantly related animals, which is expressed, for example, in different welcoming rituals.

Bachelor group of African elephants in Tsavo East National Park in Kenya.

The bulls show a different behavior towards the cows. They leave the maternal herd around the age of nine. Initially, they join other family associations, usually near their place of birth. Sometimes they change the bandage more often. Later in the fully sexually mature age, the individual individuals form bachelor groups. Here they mostly stay during the sexually inactive phases. During the musth , the sexually active time, they wander around as solitary animals in search of cows ready to mate.

In principle, the African elephant is not territorial. The family groups use activity spaces that can vary in size depending on the habitat. In more humid landscapes such as in Lake Manyara National Park in Tanzania, sizes of 15 to 52 km² were determined, in drier landscapes such as in Tsavo East and Tsavo West National Park in Kenya the areas of activity reach dimensions of 350 to 1580 km², in the Kruger -National Park in South Africa between 1600 and 4000 km². Animals have the largest action areas in desert-like landscapes such as in the Zakouma National Park in Chad with an area of ​​almost 4700 km², in the Etosha National Park in Namibia with 5800 to 8900 km² or in the Kunene region, also Namibia, with up to 12,800 km². The bachelor groups of the bulls have on average larger activity areas than the family groups of the cows, but there are hardly any differences between the sexes in the absolute dimensions. Furthermore, the extent varies with the season, so that the residence areas are larger in the rainy season than in the dry season. There are different zones within the action areas, which are used in different seasons. The individual herds migrate in the annual cycle between the residence areas and cover an average of 5 to 13 km daily with shorter distances during the dry season. The populations from the Gourma region in Mali cover around 3900 km annually. The herds migrate in an annual circular cycle that takes them to the watering holes in the north in the dry season. With the approaching rainy season, the migrations are directed south again, which can be explained by the larger amount of food available. In southern Africa, the herds change annually between the Kruger National Park and the neighboring Sabi Sand Reserve . In both protected areas, they use core zones relatively close to the water, although the animals are rarely more than 3 km from the nearest water point. Your daily hikes amount to a minimum of around 4 and a maximum of 44 km.

The greatest migration speed of the animals in Mali is around 6.1 to 6.4 km / h. On average, however, a herd usually only covers just under 0.5 km per hour. The walking speed increases in landscapes with strong human influence. On average, an animal in the low speed range consumes around 3.2 joules per kilogram of body weight over a meter covered. The energetic costs decrease continuously with higher step frequency. The net energy at rest is around 0.8 joules per kilogram of body weight. The African elephant is one of the land mammals with the lowest consumption. It is significantly below that of a mouse, which can reach 20 to 40 times the value in relation to body size. The individual animals and smaller family groups sometimes move independently of one another in the action areas, but coordinate their wanderings with one another using low-frequency resentments. The paths that emerge during the hikes are often recognizable in the vegetation over several kilometers and sometimes up to 1.3 m wide.

Agonistic behavior and communication

Welcome ritual of two African elephants
Two fighting African elephants

With a few exceptions, the coexistence of the African elephant is rather peaceful. Animals of reunited herds carry out a welcoming ritual, which is expressed by holding their heads high, snuggling trunks, inserting the trunk into the other person's mouth and urinating. Similar rituals also exist in the bachelor groups, but are less intense. Relationships with other family groups are also mostly friendly. Major conflicts often arise in areas with limited or irregularly distributed resources. Fights between individual animals rarely occur and are mostly highly ritualized. Sometimes they take place playfully with young animals, but also with cows during the defense of their herd. Dominant behavior is recognizable by an elevated head position over the shoulders, so that the corresponding individual appears larger, possibly associated with climbing a fallen tree trunk or an anthill. In contrast, inferior animals lower their heads far down. Frequent signs of irritation in the African elephant are spreading the ears and constant shaking of the head, but the former is also used by the visual enlargement of the signaling individual in imposing or threatening gestures. The gestures can be followed by a wide variety of reactions such as mock attacks, pushing or ramming. Signs of increased aggressiveness include shaking your head, throwing objects with your trunk, and digging with your tusks. Ambivalent, worried or indecisive postures indicate, among other things, a twisted trunk or swinging legs. If the entire family group is threatened, the members form a close heap with the young animals in the middle. Exceptions to these ritualized acts are bulls in the Musth , which are highly aggressive. This is where fights occur that do not follow a given pattern. By using the tusks, they sometimes have a fatal outcome. In addition, various other optical stimulus signals are available for communication with one another, which contain certain body and leg positions as well as the posture of trunk and ears. They can be both very obvious and extremely subtle, and combine in many ways. A feature of intensive social contact is the game that occurs not only in young animals, but also in adult animals up to old age and is played both together, alone and / or with a wide variety of objects.

In addition to the optical signal generators, there are other forms of communication with the African elephant. Among other things, the sense of smell is very important. It is used, for example, in inter-sexual communication such as when the bulls search for a partner during the mating season. In addition, chemical signals have an effect within the gender community, for example in the Musth, since the secretions of the temporal glands have individually slightly varying compositions. Female animals, in turn, can distinguish up to 30 family members based on their smell. Information carriers also form the various secretions of the inter- toe glands , ears and eyes as well as urine and feces from the anogenital tract. The sense of smell is also essential in the mother-young animal relationship. This is intensified to a large extent by the sense of touch , in which the trunk plays a central role.

Comparison of the spectrograms of social resentment in female (left) and male (right) African elephants
Social rumble of a female elephant, recorded by an acoustic camera , which visualizes the origin of the sounds

The acoustic communication of the African elephant is extremely complex. The animals are able to generate sounds in the range from 9 to 9000  Hz . The repertoire of sounds ranges from a low-frequency, dull rumble to a higher-frequency trumpet, screaming, croaking, snorting, barking and roaring that can also be heard by humans. Especially the social rumbling is very common. It acts as a contact call when herd members reunite within the fission-fusion network and at the same time coordinates the movements of the herd. After investigations on the family groups in Amboseli National Park, the individual animals differentiate between different individuals on the basis of the contact calls. In this way they recognize the herd members of the herd, but are also able to filter out foreign individuals. A single animal of a certain family group can thus keep around 100 individuals from up to 14 different herds in the area apart. It also remembers the calls of herd members, some of whom have died or who have emigrated, for several months and years. The distinctions are possible up to a distance of around 2.5 km from one another. Under certain circumstances, such as in the Etosha National Park , special atmospheric conditions at night carry the sounds up to 10 km, which increases the communication distance many times over. The tones in the low frequency range caused by self-sustained vibrations of the vocal cords comparable to human speech and singing, and not by neuronal controlled twitch muscle such as in the purring of the domestic cat . However, the animals can produce two different grudges: one nasal and one oral . The former is usually heard when looking for contact, the latter is important in the association. Both differ in the length of the generation path. In the case of the nasal rumble through the trunk, which functions as a resonance chamber lengthened by 3 m, this is more than twice as long as in the oral one. On average, the nasal rumble reaches about 19.7 Hz, a lower frequency than the oral one with 26.9 Hz.

Further studies show that cops also use social rumbling quite often. There are, however, deviations from the sounds made by the cows, which affect, among other things, the scope and the frequency used. Male animals often modulate in a lower range of 10 to 13 Hz than females, whose sounds reach 14 to 18 Hz. Among the bulls alone, the sounds of older and larger, i.e. more dominant individuals, are on average lower than those of younger and smaller animals. As a result, male animals transmit important information about their age and size in this way. In principle, it can be seen that bulls have a sound that is as extensive as cows. It is similar with young animals. Initially there are hardly any differences between the sexes. Only with advancing individual development do the differences between male and female animals emerge. Warning and alarm calls usually reach a higher frequency range. These include the aforementioned trumpets, snorting, croaking and similar sounds. Trumpet calls sound in a frequency range of 300 to 3000 Hz and last up to five seconds, snorting and croaking sounds only reach around 1000 Hz. The former are rather short at one second, the latter are held for up to ten seconds. Alarm calls vary depending on the type of threat. In the presence of humans or wild bees, the African elephant emits different and specific calls, which are also associated with different optical signals. To a lesser extent, low-frequency and high-frequency sounds are combined with one another. In addition, the African elephant is able to learn and imitate individual new sounds, as is known, for example, from only a few mammals, such as whales , and birds . It is assumed that this ability to learn is also connected to the complex fission-fusion social community and contributes to mutual recognition.

Comfort behavior and cognitive skills

Rubbing against a tree after bathing and pelting itself with sand will rid the elephant of vermin

The comfort behavior of the African elephant includes various baths in water, mud and dust. The animals either roll over on the ground or spray themselves with their trunks. Then they rub their bodies against living or dead trees or anthills. The procedure, which is sometimes carried out daily, not only protects against significant water loss from the body, but also serves to remove parasites . Sometimes the African elephant uses branches or sticks that are held with the trunk. Sometimes it also drives away annoying insects . In addition, an animal performs various movements with its trunk, mostly to suppress itching . The waving of the ears as part of the thermoregulation is striking .

in its behavioral repertoire, the African elephant shows a high level of unusual cognitive abilities. His long-term memory enables him to remember geographical locations, events or individuals and to wander over long distances. In addition, the animals are able to differentiate quantitatively deviating amounts. The various individuals in a family group can locate the position of spatially invisible herd members and relate them to their own. Furthermore, the African elephant shows a certain empathic feeling. It includes, among other things, the generally cooperative togetherness within a family group or a bachelor group. For example, animals help injured members, remove foreign objects from the bodies of relatives, drive away predators together or form alliances with one another. In many cases the behavior is aimed at the offspring, for example escaped young animals are brought back into the herd or motherless younger ones are protected. According to this, the African elephant is sensitive to the needs of other individuals and is able to recognize their individual needs; in addition, it is also able to anticipate and avoid certain stressful situations. This does not only refer to immediate relatives, but may also affect strange individuals. Other forms of cognitive ability are found in training young animals. For example, older cows fake a rut to help younger cows with their first sexual cycle. Studies have also shown that the African elephant can suffer trauma , the origin of which may extend into the fetal stage . These often arise from stressful situations in connection with man-made habitat loss , poaching or culling for population control. For example, animals from relocated family groups showed increased symptoms of stress up to six years after exposure. In connection with this is also the ability of the African elephant to be able to differentiate between different groups of people according to ethnicity , gender and age, which the animals then assign to certain hazard categories depending on their individual experience.

A certain interest in deceased conspecifics can be seen as a remarkable behavior in the African elephant. This is expressed through touching and playing with bones. According to experiments, the focus is particularly on tusks and skulls. In contrast to the view that is often given, this interest is not directed specifically towards family members, but generally towards representatives of their own species. However, it is possible that the animals visit places where relatives have died on their migration through their area of ​​activity. According to observations in the Samburu National Reserve, the animals in the vicinity of a dead conspecific sometimes show increased secretion flow from the temporal glands. There is also stronger social interaction. Similar behavior in dealing with the deceased is unknown in other mammals with the exception of humans. Only the chimpanzee deals with dead relatives for some time, but loses interest if the carcass decomposes more strongly.

nutrition

An African elephant in Kenya uses its trunk to reach the foliage in the treetop
African elephant eating

The African elephant is primarily a herbivore and feeds on grass , roots , leaves , twigs , tree bark , wood and various fruits . The diet therefore consists of a mixed vegetable diet, but the actual composition varies from region to region. It can consist of up to 70% grasses in open savannah landscapes and almost entirely of soft plants in closed forests. In mixed landscapes, the animals alternate between hard grass food and soft components, depending on the seasonal supply, depending on what currently offers the greatest proportion of nutrients and fibers. This leads to a seasonal change in food. In the Hwange National Park in Zimbabwe, the African elephant gets its food from 87 types of trees and shrubbery, 42 types of grass and 36 types of herbs . Animals from Amboseli National Park in Kenya feed on 29 different types of trees and bushes, 22 types of grass and 40 types of herbs. Investigations in the Kruger National Park in South Africa showed that the animals in the more humid south eat 10% grass during the dry season, but 50% in the rainy season. In the drier northern part of the national park, however, the proportion of grasses in the rainy season is similarly high, but the amount drops only slightly to 40% in the dry season. Presumably the conditions of the landscape play an important role here. In the north, mopane forests dominate , which are rich in nutrients and an important source of food, especially in the rainy season. In the south these are absent, the plant communities growing there are more species-rich, but potentially poorer in nutrients. As a result, the animals can and must switch to other food crops. In the Atherstone Collaborative Nature Reserve , also in South Africa, the diet in the late rainy season consists of around 60% grass and around 40% soft food. The softer plant components then increase in the dry season and make up 100% here at the height of the season . Even in the early rainy season, their share is still very high at more than 60%. The change from soft to hard plant food is particularly strong in the desert-like areas of southwestern Africa. Here, after rainfall or during the rainy season, the animals adapt completely to grasses as food and gradually switch back to leaf food in the dry season.

Especially in the more humid season the African elephant eats fresh growing Panicum , Hundszahn- , cattails , Zyper- and Urochloa grasses. The fruits of the Manketti and marula trees also form attractive sources of food, but the animals do not always eat fruit. Furthermore, analyzes of African elephants in the Chobe National Park in Botswana showed a sometimes highly selective approach to food selection. According to this, the animals prefer only 30% of the 27 more common tree species there in the rainy season. The proportion increases to up to 50% in the dry season. There is also a change from preferred buds and freshly sprouting leaves in the rainy season to branches, bark or parts of roots in the dry season. The African elephant often rasps bark off the trees with its tusks. Various acacias are preferred here, such as the umbrella acacia and the sweet thorn acacia , but also the marula tree. The bark of these trees is particularly thick and very fibrous. Due to its large body size and rapid metabolism , the African elephant needs a food amount of around 1 to 2% of its body weight, which corresponds to around 34 to 78 kg dry matter (or around 115 to 300 kg wet weight). An animal invests more than 60% of its daily time budget or up to 17 hours a day. The duration and frequency of food intake increase with the better quality of the plants during the rainy season. The feeding takes place with the flexible proboscis, with which an animal can precisely grasp individual food plants.

Elephants at a waterhole in Namibia

Water is extremely important for the African elephant, of which it needs up to 160 liters per day. The dependence on water can lead to less suitable and even degraded landscapes, especially in the dry season. This is most likely a reason for the sometimes long, seasonal migrations of some populations. If fresh water is not available, the animals may also drink brackish water , which can be harmful to health. The desert-dwelling representatives, as well as those of other regions, dig holes up to 1 m deep with their feet and with their proboscis in order to get to near-surface water. This sometimes happens in close proximity to fresh water. The water holes in turn attract other animals in the area. Sometimes the African elephant uses artificial water reservoirs and can destroy the water supply infrastructure created by humans. Salt licks and mineral-rich soils also play an important role. Some of these serve to neutralize toxic components in plants. In addition, the animals compensate for low-sodium vegetable food, which is mainly available in dry areas or during the dry season. According to studies in the Hwange National Park, cows use this more intensively than bulls and thus spend a larger part of their active time budget. The increased geophagia in female animals is mainly related to the higher energetic costs of carrying the offspring. The African elephant sometimes breaks up termite mounds in order to get to mineral-rich soil material.

Reproduction

Rut and mating

Secretion during the musth
African bull elephant in musth, the flow of secretions is clearly visible to the sides of the eyes and mouth

The mating season is all year round, but there are differences between the individual populations in different landscapes. Sometimes a certain seasonality can be determined with an increased number of births in the rainy season. Of around 1,030 registered births in Amboseli National Park, around 81% occurred between November and May. The seasonal restriction increases the drier a landscape is. In southern Africa the reproductive phase occurs exclusively in the rainy season.

The most striking characteristic of bulls willing to reproduce is the musth . It occurs in young bulls for the first time at around 29 years of age, and its duration increases with age. Initially, it only lasts a few days and then extends to up to four months. Entry into the Musth is individually different and does not run synchronously, but partly overlapping, which is a clear difference to the mostly seasonal rutting season of ungulates . A typical characteristic is the swelling of the temporal glands and the discharge of secretions from them. Furthermore, the eye-catching “musth walk” with a raised head and hidden chin as well as spread ears is part of it. The head is swung in a controlled manner. There is also excessive urination with up to 400 liters per day. The changed chemical composition of the urine during the musth is perceived by other males. In addition, the animals emit numerous low-frequency sounds around 14 Hz, which are known as "musth growls". The Musth is characterized by a higher sexual and aggressive activity of the bulls, combined with an increased testosterone level. This can reach a concentration of over 50 ng / ml blood, in extreme cases even over 100 ng / ml. In male animals outside the Musth as well as in females it is often below 1 ng / ml blood and usually does not exceed a value of 2 ng / ml. Bulls in the Musth are generally higher in rank. There are then often fights for dominance among Musth bulls or arguments with other male animals. Under certain circumstances, physically weaker individuals in the musth may dominate physically superior animals outside the musth.

Cows give birth to a calf about every 4.5 years. Her sexual cycle is extremely long, averaging 13 weeks. It consists of a luteal phase lasting around 9 weeks and a shorter follicular phase. Both sections are separated by a nonluteal phase that lasts 19 to 22 days. During this period there is a twofold increase in the production of luteinizing hormones , with only the second increase also causing ovulation . The function of the first increase in hormones has not yet been clarified. Due to the long sex cycle, cows that are not raising a calf are usually only receptive three to four times a year. Clear signs of a rut include caution and alertness with the head held high, as well as the "rut run" in which the female animal leaves the herd and watches the bull following it. At the height of the sexual cycle, the cow is accompanied by a mostly strong bull. Field observations in Amboseli National Park showed that cows also opt for older bulls in the Musth in more than two thirds of all cases. Reasons for the choice of older males could be primarily a secure fertilization and a higher probability of long-term survival of the offspring, but subordinate to the avoidance of intrusive persecution by younger individuals. Younger bulls try to climb cows - mostly unsuccessfully - in the phases before and after the peak of the rut. Likewise, younger cows often cannot prevail against more dominant family members during the rut.

Mating act within the herd

The long period between two sexual cycles of a cow and the short period of the rut itself mean that cows ready for reproduction are rare within a population. In the Musth, bulls use various strategies to track down cows in heat. They leave their traditional area of ​​action and hike long distances. The distances covered are significantly more extensive compared to sexually inactive animals. The bulls often join different family clans in the Musth, preferring larger herds to increase their reproductive success. In addition, they increasingly listen to the calls of the females. They are able to distinguish between the calls of unbound and bound female animals. Their own calls, the “musth rumble”, are not only attractive to cows in heat, but are also often answered by a female “family choir”. To make cows more attractive, bulls mark their hiking trails, trees and bushes with urine and droppings. They use the cows' excretions to track down rutting individuals with the help of the scent traces. With the help of the urine smell, which contains various pheromones , you can tell the individual phases of the rut apart. On the other hand, the cows also use variations in their sound to indicate which phase of the mating cycle they are currently in.

The sexual act takes place several times a day with an interval of 10 to 20 minutes and lasts 45 to 120 seconds. The cow makes grunting noises in the low frequency range that oscillate between 18 and 35 Hz. These are accompanied by screaming, trumpet calls and rumbling as well as by the discharge of feces and urine of the family members, what is known as the "mating pandemonium".

Birth and Individual Development

The gestation period is around 660 days. The mother usually gives birth to the offspring within the family group, but birth can also take place at any location. It often takes place at night. Typical signs of childbirth are kneeling, crouching or lying positions, knocking out, vaginal secretions or urinating. In addition, the interest of the other family members increases. The birth itself is relatively quick. Usually the young is still covered by the placenta , which is removed from the other animals in the herd. With the help of the other herd members, the newborn can get up in around 15 minutes. The entire birthing process is accompanied by up to 40 minutes of trumpeting, roaring and rumbling. Other young animals also show great interest.

Suckling young animal

In the majority of cases, a young is born, twins are less often born. According to studies in the Amboseli National Park, the proportion of twin births is 0.1%, whereas in the Etosha National Park it is 10%. Newborns weigh between 75 and 120 kg, in rare cases up to 165 kg, and have a shoulder height of 79 to 92 cm. This allows them to run under the mother's body. In general, the growth rate of the young is relatively high in the first five years. Already after the second year it runs differently between male and female calves. At this point, male calves are already larger than female ones. This is caused in part by a subtle difference between male and female pups. The mother usually always allows the former to suckle milk, but appears less tolerant in the latter. In addition, male calves suckle more often, but for a shorter time. Overall, this means that they consume more milk compared to the young female animals. The different amount of milk supplied by the mother animal means that she invests more energy in the male offspring. This is explained, among other things, by the faster growth of the male calves compared to the female. Regionally, there are sometimes significant differences in growth. In Amboseli National Park, young males reach a shoulder height of 200 cm by the age of ten, while females are around fourteen. The corresponding figures for the Kruger National Park are seven and nine years. Cows reach their full height at around 30 years of age, while bulls continue to gain size and weight into their 50s. As a result of this prolonged growth phase, older males exceed the females by 30% in height and sometimes by twice as much. The cause of this is a late closing of the epiphyseal plates of the tubular bones and the lack of adhesion of such joints on the vertebrae and the shoulder blade . The general growth of the young animals can, however, be strongly influenced by drought events or by young, inexperienced mother animals, especially in the early part of the suckling phase. The resulting growth retardation results in reduced height and higher mortality in adulthood.

Newborns are pink in color and have significantly more hair than adult animals. In addition, the eyesight is still poorly developed, so that newborns can find their way around with their trunk. In this phase of life there is a greater threat to the offspring from predators . It therefore mostly stays close to the mother and rarely moves more than a few meters away. The herd's protection at this point is also very high. The distance to the mother, however, increases with the age of the calf, with male young on average lingering further away from the mother than female. In general, mothers only give milk to their own offspring. The young are usually suckled for around two years, but weaning may take longer depending on the region. Often weaning is combined with the birth of the next calf. Weaned calves spend a large part of their daily activity eating, walking and resting. Playing with other calves is also very important. It consists of hunting, mutual climbing, rolling and fighting with the trunk or the head. Play activities sometimes increase with age. Here, male calves are increasingly looking for same-sex playmates from unrelated herds, while the female offspring tend to remain in the family group. In this way, the play of the female animals is more related to the family group and serves the social cohesion, but that of the male is directed outwards to members outside the family. Older female pups also take care of the younger offspring, which is referred to as "allomaternal care". In many, but not all cases, these are siblings. The sometimes noticeable gender-related differences in the behavior and development of the offspring reflect the later deviating reproductive behavior of the cows and bulls in adulthood.

Family group with young animals of different ages

The birth interval is around 3.5 to 4.5 years; if the environmental conditions are poor, the next birth can be delayed by up to five years. In landscapes with harsh conditions, such as in Kaokoland , intervals of more than nine years can occur. On average, cows give birth to their first calf between the ages of 13 and 15, but sometimes as early as around nine years. According to studies of over 830 cows in the Amboseli National Park, the first birth takes place at an average of 13.8 years, only 5% of all female animals give birth to a calf earlier. Mother animals and childbearing daughters can reproduce in family groups at the same time; in very large family groups, three female generations are sometimes involved in reproduction in the same period. From around nine years of age, males live largely independently of their families, and by around 14 years they are completely independent. At this point, they enter puberty , indicated by the first production of sperm and a combined testicle size of 650 to 700 g. A quantitative amount of sperm is only reached a few years later, which is considered sexual maturity. Since the young males have only reached about half of their full body weight at this point in time, they cannot yet enter into a mating competition with older bulls. A sexual cycle first occurs in the early 1920s. A high point of fertility generally becomes apparent between the ages of 16 and 40. However, both female and male African elephants can reproduce well into the 1960s, although the reproductive rate declines from the 1950s. The maximum age in the wild is unknown, but it is generally assumed to be 60 to 65 years for both sexes and is individually dependent on the quality of the food and the stress on the teeth. Life expectancy in undisturbed populations averages 54 years for cows and 39 years for bulls. It sinks rapidly in landscapes with high hunting pressure on the part of humans. In the Amboseli National Park it is 41 years for cows and 24 years for bulls. The corresponding values ​​in the Samburu National Reserve are only 22 and 19 years. In general, male animals are exposed to a higher mortality rate. In human captivity, animals have reached a maximum of around 55 years.

Predators, parasites and ecology

Predators and interactions with other animal species

Lions as one of the few predators of the African elephant

A fully grown African elephant has few natural enemies. Calves and young animals up to 15 years old are occasionally preyed on by lions and spotted hyenas . The influence of predators on local African elephant populations varies with the seasons. It also depends on the density of the predator population and the size of other potential prey. In the Hwange National Park in Zimbabwe, around 22% of all documented prey raids by lions include elephants, with young animals under the age of four being killed. Overall, elephants reach around 16% of the captured biomass. According to observations in Chobe National Park in Botswana, almost every fourth attack by lions on elephants is successful. These often take place at water points, and the animals killed are between four and eleven years old.

There is a rich spectrum of interactions with other animal species. Calves and young animals often play with birds, monkeys, warthogs and antelopes. In adult individuals, there is largely competition that exists in open landscapes with the two African rhinos, the hippopotamus and the Cape buffalo . The African elephant also chases lions and hyenas away from their prey or from the vicinity of family groups. It can also kill other animals such as birds or smaller mammals in a targeted manner, including when frightened. Especially young bulls in their first musth sometimes show extreme aggression against other larger mammals. Alone Hluhluwe-Imfolozi Park in South Africa introduced young bulls killed between 1991 and 2001, nearly 60 Breitmaul- and five black rhinos . Likewise, almost 50 white rhinos were killed by elephants in the Pilanesberg National Park in the 1990s. This is mostly due to the lack of higher ranking older bulls, through whose presence the musth of the young bulls can be controlled and directed.

Furthermore, the African elephant reacts to the humming noise of wild and aggressive bees by fleeing. This implies that the animals recognize bees and relate their flight noise to negative events from the past. An individual or a related family member may have been stung, but the experience may also be from social learning.

Parasites

Numerous parasites of the African elephant have been documented. The outer ones include more than 20 types of ticks , as well as lice such as Haematomyzus or ear mites such as Loxonaetus and Otonaetus . In addition, there are a wide variety of flies such as bot flies and stomach flies . Particularly important are Pharyngobolus , Pltycobboldia , Ruttenia and Cobboldia . Internal parasites include over 30 known species, including nematodes as Grammocephalus and flukes as Protofasciola . Investigations on faecal samples from the Okavango Delta show that the infestation is particularly high in the rainy season. In addition, the family clans are more affected than the bachelor groups.

Anthrax can have a greater impact on local populations . In Etosha National Park, a total of three epidemics with over 300 dead African elephants were registered in the 1980s . Encephalomyocarditis , which is caused by the picornavirus and transmitted by rodents, is also fatal in individual cases . The disease killed more than 60 African elephants, most of them bulls, during a major outbreak in 1994 in Kruger National Park. The floppy or flaccid trunk syndrome is largely unexplored . The nerves of the trunk degenerate, which can lead to complete paralysis of the organ. For affected individuals, this can be fatal as food intake is no longer possible. The disease may be caused by plant toxins.

Ecological importance

Elephant dung; the woody components of the food can be recognized.

The African elephant has a very strong impact on its environment. The debarking or kinking of trees, mainly or often by bulls, damages woodland, for example. With a correspondingly high density of a local population, larger areas of their tree population can be robbed and turn into grassland. Associated with this is a decline in the biodiversity of the local flora and fauna. Such changes are documented, among others, from the Murchison Falls National Park in Uganda, the Tsavo East National Park in Kenya or the Kruger National Park in South Africa. Little data is currently available on the long-term effects. Both elephants and trees are long-lived creatures, and their interactions have so far been little studied. Numerous external factors such as hunting or loss of habitat in individual regions also interfere with the investigations. There are also other, natural occurrences like fire. Bush fires, for example, can dissolve dense stands of trees, while elephants continue to keep the landscapes open in the following. Various models have therefore been developed for the influence of the African elephant, ranging from equilibrium through stable cyclical processes to repetitive cyclical events.

The opening up of wooded landscapes, however, also means that these areas become accessible to other animal species of the savannah, such as rhinos, and that these also use the new refuges. On the other hand, an increasing local population of the African elephant can help to transform artificially altered landscapes back into a more natural state. On a smaller scale, the cavities created when debarking trees or breaking branches serve as places of retreat for small vertebrates. This can sometimes have a significant influence on their populations, as has been demonstrated, for example, in geckos of the genus Lygodactylus . Furthermore, the paths laid out by the African elephant during its migrations are of great importance for other animal species.

The African elephant also has a larger share in the spread of plant seeds that it excretes with its dung . The more common seeds include those from acacia, but also from the marula tree, from mangoes and melons, and from the genus Balanites . The latter, which is to dry forest trees from the family of Lignum-vitae is, benefited significantly from the African elephant as it According to analyzes in Uganda at slightly more than half of the seeds after passage through the gastrointestinal tract for germination occurs. For seeds that did not pass through the digestive system, it is less than 1%. The passage through the gastrointestinal tract takes an average of 24 to 36 hours. The migration behavior of the animals enables the seeds to be transported over distances of around 2.5 km, in extreme cases up to 65 km have been observed.

Systematics

Internal systematics of today's elephants according to Meyer et al. 2017
 Elephantidae  
  Loxodonta  

 Loxodonta africana


   

 Loxodonta cyclotis



   

 Elephas



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The African elephant is a species from the genus of the African elephants ( Loxodonta ). The genus also includes the forest elephant ( Loxodonta cyclotis ) as a further representative. Loxodonta , the sister group of the genus Elephas with the Asian elephant ( Elephas maximus ) as the only member. Both genera in turn belong to the family of elephants (Elephantidae) in the order of mammoths (Proboscidea). The proboscis form a very old group of originally African animals. Its origins go back around 60 million years. In the course of their tribal history, they proved to be very diverse with adaptations to numerous biotopes and climatic regions. The trunk animals also colonized large parts of Eurasia and America . Compared to the long tribal history of the proboscis, the elephants can be viewed as a relatively young line of development. The first representatives can be found in the late Miocene . Precursors of today's forms appeared in fossil form around 7 million years ago. This roughly agrees with molecular genetic studies, according to which the African and Asian elephants separated from each other 7.6 million years ago. The two current members of the African elephant differentiated themselves from 5.6 to 2.6 million years ago.

In general, no subspecies of the African elephant are distinguished today. This looked different in the research history past. Up until the 1990s, all elephants on the African continent were considered to belong to a species within which up to six subspecies were recognized. These six subspecies of the African elephant in the broader sense were divided into two different groups:

  • L. a. africana division
  • L. a. africana ( Blumenbach , 1797); southern africa
  • L. a. knochenhaueri ( Matschie , 1900); eastern africa
  • L. a. orleansi ( Lydekker , 1906); northeastern Africa
  • L. a. oxyotis ( Matschie , 1900); northern africa
  • L. a. cyclotis division

In the course of the 20th century, the forest elephant only held the position of a forest variant of the African elephant. Only various genetic and anatomical studies in the transition from the 20th to the 21st century led to the recognition of the forest elephant as an independent species. Both representatives of the genus Loxodonta hybridize in the contact area of ​​their distribution areas. The division into the other subspecies is largely doubted today, especially since genetic studies cannot understand this. The African elephant is therefore to be regarded as monotypical , even if there are numerous variations in body size and tusks in the entire distribution area.

Tribal history

The genus Loxodonta first appeared in Africa in the Upper Miocene around 7 million years ago. It developed from Primelephas , the oldest representative of the real elephants in terms of phylogeny. However, some paleontologists also assume an origin of Stegotetrabelodon . Several extinct species are known, all of which are confined to the African continent. The fossil forms include Loxodonta cookei, the oldest form to date , as well as Loxodonta adaurora , Loxodonta exoptata and Loxodonta atlantica . Today's African elephant can be grasped for the first time in the transition from the Lower to the Middle Pleistocene , but its predecessor is unknown. The form Loxodonta atlantica , which occurs around the same time in northern and southern Africa, is considered too modern to be in the direct line of the African elephant. Likewise, the only slightly older species Loxodonta adaurora can be ruled out, which is very similar to the African elephant in its skull structure, but differs markedly in its teeth structure. Confirmed finds of the African elephant from this period include the Awash in the Afar triangle in Ethiopia. It is possible that some teeth and a skull of a young animal from Angamma-Yayo in Chad extend the temporal occurrence of the species into the beginning of the Lower Pleistocene. Other early finds come from the Kibish Formation on the Omo , also Ethiopia, and from Elandsfontein in South Africa. However, they already date to the late Middle Pleistocene. Significantly more recent finds were also documented from the latter site.

Research history

Johann Friedrich Blumenbach

Knowledge of the African elephant in the western world goes back to ancient times . These then increased especially at the time of the colonization of Africa. The term "Ivory Coast" for part of western Africa refers to the large quantities of ivory that was shipped from there to Europe. A living African elephant from this region reached the court of King Louis XIV in 1668, he lived in the gardens of Versailles until his death in 1681. The skeleton of the animal was used by Georges-Louis Leclerc de Buffon and Louis Jean , among others -Marie Daubenton 1769 for a detailed description. Even Georges Cuvier used it in 1806 together with some specimens from southern Africa for an in-depth processing. For the animal from West Africa, the remains of which are still in the Muséum national d'histoire naturelle in Paris, an assignment to the forest elephant is likely.

In 1754, Carl von Linné only led the Asian elephant under the name Elephas indicus . In the tenth edition of his important work Systema naturae from 1758 he summarized both the Asian and the African elephant and named both with Elephas maximus . Almost forty years later, in 1797, the German natural scientist Johann Friedrich Blumenbach undertook a formal separation of the two elephant species in his "Handbuch der Naturgeschichte". He referred the African elephant to the species Elephas africanus and justified this step with the rhombic lamellar structure of the molars, in contrast to the Asian elephant. Blumenbach is therefore considered to be the first scientific descriptor of the African elephant. The different tooth structure then prompted Frédéric Cuvier to define the genus Loxodonta in 1825 ; he introduced the term within a monograph on the natural history of mammals, but dispensed with Latinization . It was not until 1827 that the generic name Loxodonta was correctly reproduced by an unnamed author in a review of the corresponding work, so that it is now considered the first name of the genus.

The finds with which Blumenbach set up his new species, Elephas africanus , was in the collection of the University of Göttingen and is probably lost today. It can be assumed that it originally came from southern Africa - as is often the case with early research material. Hermann Pohle therefore established the area around the Oranje River as a type area in 1926 . From the same region, Georges Cuvier had already assigned the name Elephas capensis to the African elephant one year after Blumenbach and independently of this . Today the name is synonymous with Loxodonta africana . Other names were introduced during the 19th century. However, there was never a stronger division of the species into different subspecies during this period. Only Matschie led in 1900 a number of subspecies, including the forest elephant. Only a little later in 1907 Richard Lydekker tried to structure the species more strongly based on the shape of the ears. The procedure was later criticized because of the high variability of the feature. With the exception of the forest elephant, the subspecies postulated by Matschie and Lydekker are no longer recognized as a separate form.

African elephant and human

Local art and culture

Depiction of an African elephant in the Tadrart Acacus

The African elephant has been of great importance to the local population as a source of food and raw materials for thousands of years. In addition to the meat, the skin and ivory of the tusks were mainly used. As one of the most impressive animals in the African landscapes, it has also been depicted in local art across the continent. Outstanding here is the wall art of the San in southern Africa. In the community of Cederberg in South Africa alone there are around 200 rock overhangs with elephant images. Some of them are intensely red. The very realistic reproduction of the animals can be seen as remarkable, in terms of both body proportions and lifestyle. The pictures show, among other things, family groups hiking or eating.

Another important center of African rock art can be found in the Sahara in northern Africa. The oldest rock art was created here as early as the Bubalus period , which covers the transition from the end of the Pleistocene to the beginning of the Holocene and is characterized by depictions with curved lines and distorted proportions of the so-called "Tazina school". The regions of the Messak or the Tadrart Acacus of Libya can be named as outstanding . The former includes drawings of giant animals with tiny people holding curved objects, probably boomerangs , to their trunks, while the latter stands out with a depiction of a herd, the rearmost individual of which appears to have been defecated. More recent are the animal motifs from Wadi al-Ajal in southwestern Libya, which date from about 6300 to 1000 BC. Belong to the cattle age . The style of this phase is characterized by sub-naturalistic, punched images. In the Wadi al-Ajal, a good 3.1% of all depictions depict the African elephant, which numerically corresponds to 37 individuals. It should be noted here an engraving of a bull in the musth. In southern Morocco in the Akka region , the African elephant makes up around 2.6% of all depictions of the bubalus and cattle ages. Very often there are groups in which the different sizes of the individuals are reminiscent of mother-young animal associations. Occasionally representations of elephants appear together with those of humans. Subsequent motifs of the African elephant appear again and again, for example in Morocco. In addition, elephant representations are also available from western and eastern Africa.

taming

Hannibal crosses the Alps with war elephants, fresco in the Palazzo del Campidoglio, Rome, around 1510

Unlike the Asian elephant, the African elephant has probably rarely been domesticated. However, attempts go back to the Ptolemaic times of the 4th and 3rd centuries BC. BC back. Ptolemy II laid the foundation for taming around 270 BC. BC, who needed new animals for his war elephants , which originally came from Asia . He procured these from the region south of Egypt, which resulted in the founding of several cities along the Red Sea , such as Ptolemais Theron and Berenike . The Ptolemies used this to ship the captured animals to Egypt. The effort ultimately culminated in the Battle of Raphia in 217 BC. BC, in which African and Asian elephants faced each other (it is therefore also called "Battle of the Elephants"). In research, however, it is controversial whether it was the African elephant or the smaller forest elephant. This debate is sustained by ancient written documents that attribute the Ptolemy elephants to be smaller than the Asian elephants, that they did not have towers and that they were inferior to the opposing animals. DNA analyzes on elephants from Eritrea, the likely region of origin of the animals of the Ptolemies, revealed a clear relationship with the larger African elephant in 2013. There is a similar discussion about the animals that Hannibal found during the Second Punic War around 218 BC. BC when he crossed the Alps .

Threat and protection

Elephants in South Africa

Ivory hunting and poaching have contributed to the dwindling population of the African elephant in many parts of Africa. Further threats can be found in the habitat losses caused by the increasing human population and the resulting conflicts with local residents. In individual areas of western, eastern and southern Africa it died out completely, for example in Gambia in 1913, in Swaziland in 1920, in Burundi in the 1970s and in Mauritania in the 1980s. The recent losses include Sierra Leone in 2010, while the species may also have disappeared from Guinea-Bissau and Senegal . In South Africa , between the 19th and 20th centuries, the African elephant was on the verge of extinction. The establishment of individual protected areas such as the Addo Elephant National Park in 1931 or the Kruger National Park in 1926 led to an increase in the population to currently around 18,000 individuals.

After the ivory trade was banned worldwide in 1989, there was a steady increase in stocks in East Africa and especially in the states of southern Africa. The decision was controversial and since 1997 there have been considerations to partially reopen trade and allow it to be controlled. This also led to the establishment of the MIKE program ( Monitoring of Illegal Killing of Elephants , in German about "observation of the illegal killing of elephants"). The proceeds of the trade in stock or stored tusks, which has been permitted again since 2008, has been used since then in conservation programs for the African elephant. The IUCN , which the African elephant from 1996 to 2003 as a "high risk" ( endangered had listed) species, classified him in 2004 in the lower rank "endangered" ( vulnerable ) back, which he is still performed today. Simultaneously with the worldwide trade ban, the species was placed on Appendix I of the CITES Agreement. In the meantime, the African elephant has been downgraded to Appendix II for Botswana, Namibia, Zimbabwe and South Africa, and ivory stocks from these countries are being auctioned legally. Since then, there has been an increase in poaching in numerous countries of origin.

The African elephant is found in a total of 37 countries in sub-Saharan Africa and is present in numerous protected areas. According to assumptions, the total range of the species is 70% or more outside of protected areas. Other protective measures include habitat management, which is necessary as overgrowth of local populations can negatively affect the regional landscape, other animal species, or the elephants themselves. Large-scale slaughtering ( culling ) of extremely dense stocks is no longer carried out today, but rather individual herds are relocated. To strengthen local agriculture and to protect the fields from being plundered by the African elephants, fences with beehives are being built to exploit the animals' natural fear of insects. Other measures include burning briquetted hot chilli peppers , making loud noises or bright lights, which can locally change the behavior of the African elephant. In some countries like South Africa, Tanzania or Zambia, sport hunting is allowed. The income gained from this flow back partially into the local communities and supports the tolerance of the population towards the African elephant and reduces conflict situations. In Botswana, which has the largest population with around 130,000 elephants, hunting was banned in 2014, with negative effects for communities and an increase in poaching. In addition, there are individual agreements between neighboring states for joint protective measures for highly migratory and thus cross-border groups of African elephants. Nonetheless, poaching remains a general problem. In 2009 it was estimated that 38,000 African elephants were poached annually for the ivory trade.

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This version was added to the list of articles worth reading on March 11, 2006 .