African forest elephants occur in central and western Africa. They can be found in northern Congo, southwestern Central African Republic, the southeast coast of Gabon, southern Ghana, and in Cote D'Ivoire. Many of these populations are isolated from each other currently. (Dudley, et al., 1992; Merz, 1986a; Merz, 1986b; Tangley, 1997; White, et al., 1993)
African forest elephants are most populous in central and western Africa. Areas of high density are characterized by lowland tropical rainforests, semi-evergreen and semi-deciduous tropical rainforests, and swamps. Elephants change habitats seasonally, inhabiting swampy areas during the dry season, then moving back to lowland rainforest areas in the wet season. African forest elephants seek refuge in rainforest preserves because they are persecuted by local human populations. They are illegallly hunted for ivory and killed for crop-raiding. (Dudley, et al., 1992; Fay and Agnagna, 1991; Merz, 1986a; Merz, 1986b; Tangley, 1997; White, et al., 1993)
African forest elephants are smaller than their close relatives, savanna elephants (Loxodonta africana). Male shoulder height is greater than female shoulder height and height increases with age. Mean shoulder height is between 144 and 155 cm (range 69 to 216 cm). Hind footprint length is helpful in determining age, mean hind footprint length is 24.7 and the median is 25.8 cm (range 12.5 to 35.3 cm). Boli width, or feces, is also a helpful factor in determining body size and, therefore, age. Average boli width is 10 cm (range 4 to 16 cm). Measurements of wild animals indicate that African forest elephants stop growing at about 10 to 12 years of age, younger than their savannah elephant relatives. Furthermore, growth rates are lower than savannah elephants. African forest elephants also have straighter and thinner tusks that hang more vertically to assist in passage through dense forests. They have rounded ears, unlike savanna elephants that have overhanging flaps along the upper edge of the ear. There are also differences in skull morphology between the two species: African forest elephants have less pneumatization, the formation of air cells or cavities in tissue, in their skulls than savanna elephants. African forest elephants have 4 toes on their front feet and 3 on the hind feet, African savanna elephants have 5 toes on their front feet and 4 on the hind feet, although hybrids occur. (International Elephant Foundation, 2008; Kingdon, 1979; Morgan and Lee, 2003; Roca, et al., 2001)
Elephants have the largest brain of any land animal. The brain is located in the back of the skull away from the forehead. Their forehead holds a sponge-like bone which compensates for the heavy weight of the trunk. The trunk is more sensitive than human fingers and is used to signal, trumpet, eat, bathe, dust, smell, and in defense. Trunks are also used in respiration, especially when elephants swim. They can hold their trunks above the water and breathe through them. The large ears of African forest elephants help them regulate temperature because they have few sweat glands. They cool themselves by making a fanning motion with their ears and pump blood into the ears to help dissipate body heat. Elephants have sensitive skin and can be prone to sunburn, especially when young. Their wrinkled skin also helps in keeping them cool because water is trapped in the cracks and crevices, which then evaporates. Elephants can lose 75% of their body heat using this method of cooling. The large feet and thick fat pads on the feet act as shock absorbers to help evenly distribute the pressure of their large body mass. Their feet are sensitive and can pick up vibrations through the ground, including thunder and elephant calls from up to 10 miles away. (Kingdon, 1979; Morgan and Lee, 2003; Roca, et al., 2001)
African forest elephants are polygynous. Males compete for access to estrus females and older, larger, more dominant males generally mate with more females. Males experience "musth," a hormonal state marked by increased aggression. During this period, the male secretes a fluid from the temporal gland between the eye and ear. Musth begins at 15 and 25 years old. There is a positive correlation between age and period of musth in male elephants. Younger males usually experience musth for a shorter period of time while older males usually experience it for a longer period of time. There are several behaviors that male elephants do while in musth. They have a more erect walk - their head held high and their tusks tucked inward. They may also rub the side of their heads on trees or bushes in order to spread their musth scent. They may wave or flap their ears which would blow the musth smell towards other elephants. At the same time the ear wave is performed, a musth rumble may be transitted. A musth rumble is a distinct set of low frequency calls they may be as low as 14 Hz. Younger males perform these calls less frequently than older males and females usually respond with their own calls. The last behavior associated with musth is urination. Males dribble urine slowly and allow it to spray onto the insides of their hind legs. All of these behaviors serve to advertise their musth state to receptive females and competing males. Females go through four stages of estrous behavior. The first stage is wariness, the next stage is the estrous walk, where the female leaves her group, walking away with her head held high and turned to the side watching the male follow her from behind. The third stage is the chase. The fourth and final stage of estrous behavior is consortship. A male and female elephant interact physically during this stage. The male wards off any other males as the female gets closer to her partner. (International Elephant Foundation, 2008; Sukumar, 2003)
Because African forest elephants are a recently recognized species, there is little specific information available on their reproduction. Information reported here is for the previously recognized inclusive species - African elephants. Younger males go into musth during the dry season, but older males go into musth during the wet season, when more females go into estrous. Estrous lasts for about 2 days and occurs approximately every 15 weeks. Gestation lasts 20 to 22 months, after which a single young is born, although twins occur rarely. Females nurse their young for up to 6.5 years, although young begin to include vegetation in their diet in their first year of life. Male young may nurse more than female young and allosuckling occasionally occurs - where other female members of the group nurse another female's young. Age at sexual maturity varies with climate, habitat, and diet. Wild female elephants usually reach sexual maturity between 11 and 14 years old (range 9 to 22 years). Males also reach sexual maturity at 11 to 14 years, although males typically do not successfully reproduce until they are older, more experienced, and larger. (International Elephant Foundation, 2008; Sukumar, 2003)
Females in African forest elephant groups all contribute to the care of young. Mothers nurse and care for their young for extended periods of time, from birth to about 8 years old. Very young calves stay within about 5 meters of their mother at all times. When a calf is born, it can stand on its own shortly after birth. Calves are nursed exclusively for 3 months and weaned at 78 months old. Calves spend many years learning to navigate their complex environments and find food. (International Elephant Foundation, 2008; Sukumar, 2003)
There are no reports of lifespan in Loxodonta africana) suggests lifespans of 65 to 70 years old in the wild. A living 53 year old elephant is the oldest recorded age for captive African elephants, but expected lifespan in captivity is closer to 33. (de Magalhaes, et al., 2005; Wiese and Willis, 2004). Data from the previously recognized inclusive species, African elephants (
African forest elephants travel in smaller groups than other elephant species. A typical group size consists of 2 to 8 individuals. The average family unit is 3 to 5 individuals, usually made up of female relatives. Most family groups are a mother and several of her offspring, or several females and their offspring. Female offspring are philopatric, male offspring disperse at maturity. Unlike African savanna elephants (Loxodonta africana), African forest elephants ( ) do not usually interact with other family groups. Male African forest elephants tend to be solitary and only associate with other elephants during the mating season. Males have a dominance hierarchy based on size. (Morgan and Lee, 2003; Sukumar, 2003; Tangley, 1997)
Home range size of African forest elephants is difficult to determine due to their dense forest habitat. Home range size will vary with regional characteristics, such as proximity to water and quality of food resources. (Sukumar, 2003)
African forest elephants, along with other elephants, communicate among widely dispersed social groups through low-frequency sounds (5 Hz). Because (Dudley, et al., 1992; Kingdon, 1979; McComb, et al., 2003; Morgan and Lee, 2003; Roca, et al., 2001; Tangley, 1997)is newly recognized, there is no literature on communication and perception in this species. However, it was previously recognized that African elephants are capable of recognizing a call from a family member up to a distance of 2.5 km, but can better recognize it at 1.0 to 1.5 km. Given the difference in habitat structure (savannah vs. forest), it may be expected that detection distances are shorter in African forest elephants. Hearing and smell are the two most important senses for these mammals. They can hear vibrations through the ground and can use their sense of smell to detect food sources. Like other elephants, though, African elephants have good eyesight and extremely sensitive tactile perception through their trunks and skin. Trunks are used extensively to manipulate objects and for information gathering. Elephants touch their trunk to an object, then insert the trunk into the mouth, where the chemical cues are picked up in the roof of the mouth.
African forest elephants are herbivorous, their diet is composed mainly of fruit, leaves, bark, and twigs of rainforest trees. They consume a wide variety of fruit, including Antidesma vogelianum, Omphalocarpum species, Duboscia macrocarpa, Swartzia fistuloides, and Klainedoxa gabonensis. Tree species eaten include legumes such as Piptadeniastrum africanum, Petersianthus macrocarpus, and Pentaclethra eetveldeana. Diets vary regionally with available trees and fruits. African forest elephants supplement their herbivorous diet with minerals that they get by eating soil. (Eggert, et al., 2003; Tangley, 1997; White, et al., 1993)
Humans are the greatest threats to African forest elephants. They have been extensively hunted for their ivory, which may be why many African forest elephants travel and feed at night. They are also persecuted by farmers for the damage they do to crops. Crop destruction is more often the result of foraging by rats (Nesomyidae), porcupines (Hystricidae), monkeys (Cercopithecus), and river hogs (Potamochoerus porcus), but many any crop destruction on elephants. Very young African forest elephants that somehow are separated from their family group or are ill may be preyed on by large carnivores, such as lions (Panthera leo) or hyenas (Hyaenidae), although these predators are rare in African forest elephant habitats. (Dudley, et al., 1992; Tangley, 1997; Weber, et al., 2001)
African forest elephants are important dispersers of seeds through their consumption of fruit. Forest dates (Balanites wilsoniana) are considered elephant dependent because the seed germinates more successfully after passing through an elephant. The decline of elephant populations in West African rain forests in Upper Guinea is correlated with forest date population declines. African forest elephants are also responsible for creating and maintaining large clearings in the middle of rainforests in areas where they extract mineral salts from the soil or tear down vegetation as they eat and travel. These forest openings strongly affect forest tree regeneration and increase forest tree diversity, affecting many other organisms in that area. The paths and holes in the ground that they make as they travel are used by other smaller animals for shelter or become a source of drinking water. Because of their large impact on forest composition, African forest elephants might be considered a keystone species.
Because African forest elephants were long-considered a subspecies of the inclusive African elephant species, Loxodonta africana, parasites for both species have not been sorted out. However, parasites recognized the inclusive species include: 2 species of trematodes, 32 species of nematodes, 21 species of ticks, 1 louse species, botflies, protozoans (Babesia), and blood-sucking flies in the family Anthomyidae. (Eggert, et al., 2003; Laursen and Bekoff, 1978; Sukumar, 2003)
African forest elephants have affected human populations in a positive way because they provide humans with ivory, hide, and meat. Different cultures have historically used elephant ivory for different purposes and it is viewed as a symbol of wealth and luxury. Ivory has been used for knife handles, combs, toys, piano keys, billiard balls, furniture, and artwork. Portuguese in the late fifteenth century exported 100 to 120 tons of ivory average every year from western Africa. In the late nineteenth century, the Congo exported 352 tons of ivory per year. Elephant ivory is no longer legally traded and poaching for ivory is a major threat to elephant populations. African forest elephants are important members of native ecosystems, impacting forest regeneration and composition. (Sukumar, 2003)
African elephants have been known to cause damage to crops. They typically are attracted to sugarcane (Saccharum officianarum) and cereals like corn (Zea mays), sorghum (Sorghum vulgare), and wheat (Triticum vulgare). They are also attracted to many different types of millet such as finger millet (Eleusine coracana), little millet (Panicum miliare), and bulrush millet (Pennisetum typhoides). Elephants are attracted to different fruits and vegetables such as mango, banana, orange, melon, jackfruit, potato, tomato, carrot, spinach, and pumpkin. Much of the crop damage is caused by elephants trampling over crops and is only sometimes due to the eating of crops. Other crops often affected by African forest elephants include oil palm (Elaeis guineensis), oranges (Citrus sinensis), and cacao (Theobroma cacao). Elephants occasionally kill people. These are usually chance encounters where people accidentally or intentionally become too close to an elephant, causing it to feel threatened. (Sukumar, 2003)
The IUCN Red List considers African forest elephants a subspeces of African elephants, which they consider near threatened. CITES lists African elephants under appendices I and II. Appendix I states that the animal is threatened to extinction and trade of the animal is only allowed under certain circumstances. Appendix II states that the animal is not necessarily threatened to extinction, but trade of the animal is closely monitored.
Until 2001, African forest elephants were considered a smaller, forest-dwelling subspecies of the inclusive African elephant species (Loxodonta africana). However, clear genetic differences suggest that African forest elephants are quite distinct and deserved species status. Morphological and behavioral differences recognized at the subspecies level also support this distinction. (Roca, et al., 2001; Vogel, 2001)
Tanya Dewey (editor), Animal Diversity Web.
Tara Connor (author), Radford University, Karen Francl (editor, instructor), Radford University.
living in sub-Saharan Africa (south of 30 degrees north) and Madagascar.
uses sound to communicate
having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.
uses smells or other chemicals to communicate
helpers provide assistance in raising young that are not their own
ranking system or pecking order among members of a long-term social group, where dominance status affects access to resources or mates
animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.
an animal that mainly eats leaves.
A substance that provides both nutrients and energy to a living thing.
forest biomes are dominated by trees, otherwise forest biomes can vary widely in amount of precipitation and seasonality.
an animal that mainly eats fruit
An animal that eats mainly plants or parts of plants.
offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).
a species whose presence or absence strongly affects populations of other species in that area such that the extirpation of the keystone species in an area will result in the ultimate extirpation of many more species in that area (Example: sea otter).
having the capacity to move from one place to another.
the area in which the animal is naturally found, the region in which it is endemic.
active during the night
generally wanders from place to place, usually within a well-defined range.
chemicals released into air or water that are detected by and responded to by other animals of the same species
having more than one female as a mate at one time
rainforests, both temperate and tropical, are dominated by trees often forming a closed canopy with little light reaching the ground. Epiphytes and climbing plants are also abundant. Precipitation is typically not limiting, but may be somewhat seasonal.
Referring to something living or located adjacent to a waterbody (usually, but not always, a river or stream).
scrub forests develop in areas that experience dry seasons.
reproduction that includes combining the genetic contribution of two individuals, a male and a female
one of the sexes (usually males) has special physical structures used in courting the other sex or fighting the same sex. For example: antlers, elongated tails, special spurs.
associates with others of its species; forms social groups.
a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.
uses touch to communicate
Living on the ground.
The term is used in the 1994 IUCN Red List of Threatened Animals to refer collectively to species categorized as Endangered (E), Vulnerable (V), Rare (R), Indeterminate (I), or Insufficiently Known (K) and in the 1996 IUCN Red List of Threatened Animals to refer collectively to species categorized as Critically Endangered (CR), Endangered (EN), or Vulnerable (VU).
the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.
movements of a hard surface that are produced by animals as signals to others
uses sight to communicate
reproduction in which fertilization and development take place within the female body and the developing embryo derives nourishment from the female.
breeding takes place throughout the year
young are relatively well-developed when born
Berger, L. 2001. "Opinion: How Do You Miss a Whole Elephant Species?" (On-line). National Geographic. Accessed September 23, 2008 at http://news.nationalgeographic.com/news/2001/12/1217_leeelephant.html.
Douglas-Hamilton, O. 2008. "Africa's Elephants: Can They Survive?" (On-line). National Geographic. Accessed September 23, 2008 at http://ngm.nationalgeographic.com/2008/09/africa-elephant/douglas-hamilton-text/12.
Dudley, J., A. Mensah-Ntiamoah, D. Kpelle. 1992. Forest elephants in a rainforest fragment: preliminary findings from a wildlife conservation project in southern Ghana. African Journal of Ecology, 30/20: 116-126.
Eggert, L., J. Eggert, D. Woodruff. 2003. Estimating population sizes for elusive animals: the forest elephants of Kakum National Park, Ghana. Molecular Ecology, 12: 1389-1402.
Fay, M., M. Agnagna. 1991. A population survey of forest elephants (Loxodonta africana cyclotis) in northern Congo. African Journal of Ecology, 29/3: 177-187.
International Elephant Foundation, 2008. "Loxodonta" (On-line). International Elephant Foundation. Accessed January 02, 2009 at http://www.elephantconservation.org/loxodonta.php.
Kingdon, J. 1979. East African Mammals: Large Mammals. Chicago: University of Chicago Press. Accessed November 06, 2008 at http://books.google.com/books?hl=en&lr=&id=xdH_Gi92-fAC&oi=fnd&pg=PA1&dq=loxondonta+cyclotis+reproduction&ots=6l08Vsiceu&sig=Fuy-_JFSeH2vGLmMgqIAgt3jCRg#PPP1,M1.
Laursen, L., M. Bekoff. 1978. Loxodonta africana. Mammalian Species, 92: 1-8.
Macdonald, D., S. Norris. 2001. Encyclopedia of Mammals. New York, NY: Facts on File Inc..
McComb, K., D. Reby, L. Baker, C. Moss, S. Sayialel. 2003. Long-distance communication of acoustic cues to social identity in African elephants. Animal Behaviour, 65/2: 317-329.
Merz, G. 1986. Counting elephants (Loxodonta africana cyclotis) in tropical rain forests with particular reference to the Tai National Park, Ivory Coast. African Journal of Ecology, 24: 61-68.
Merz, G. 1986. Movement patterns and group size of the African forest elephant Loxodonta africana cyclotis in the Tai National Park, Ivory Coast.. African Journal of Ecology, 24/2: 133-136.
Morgan, B., P. Lee. 2003. Forest elephant (Loxodonta africana cyclotis) stature in the Reserve de Faune du Petit Loango, Gabon. Journal of Zoology, 259: 337-344.
Roca, A., N. Georgiadis, J. Pecon-Slattery, S. O'Brien. 2001. Genetic Evidence for Two Species of Elephant in Africa. Science, 293: 1473 - 1477.
Sukumar, R. 2003. The Living Elephants: Evolutionary Ecology, Behavior, and Conservation. New York, New York: Oxford University Press.
Tangley, L. 1997. In Search of Africa's Forgotten Forest Elephant. Science, 275: 1417 - 1419.
Vogel, G. 2001. African Elephant Species Splits in Two. Science, 293: 1414.
Walsh, P., L. White. 1999. What It Will Take to Monitor Forest Elephant Populations. Conservation Biology, 13: 1194-1202.
Weber, W., L. White, A. Vedder. 2001. African Rain Forest Ecology & Conservation. Pp. 207-213 in A Turkalo, J Fay, eds. Forest Elephant Behavior and Ecology. New Haven and London: Yale University Press. Accessed October 09, 2008 at http://books.google.com/books?id=iNwfN5ASigUC&pg=PA207&dq=african+forest+elephant&sig=ACfU3U2WknR4HRncZd6lRBLtwCRxOrCG-A#PPT1,M1.
White, L., C. Tutin, M. Fernandez. 1993. Group composition and diet of forest elephants, Loxodonta africana cyclotis Matschie 1900, in the Lope Reserve, Gabon. African Journal of Ecology, 31: 181-199.
Wiese, R., K. Willis. 2004. Calculation of longevity and life expectancy in captive elephants. Zoo Biology, 23/4: 365-373. Accessed December 11, 2008 at http://www3.interscience.wiley.com/journal/109580436/abstract?CRETRY=1&SRETRY=0.
de Magalhaes, J., J. Costa, O. Toussaint. 2005. "HAGR: the Human Ageing Genomic Resources." (On-line). An Age entry for Loxodonta africana. Accessed December 11, 2008 at http://genomics.senescence.info/species/entry.php?species=Loxodonta_africana.