TheAfrican forest elephant (Loxodonta cyclotis) is anelephant species native to humid tropical forests inWest Africa and theCongo Basin. It was firstdescribed in 1900. With an average shoulder height of 2.16 m (7 ft 1 in), it is the smallest of the three living elephants. Both sexes have straight, down-pointing tusks, which begin to grow at the age of 1–3 years.
The African forest elephant lives in highly sociable family groups of up to 20 individuals comprising adult cows, their daughters and sons. When young bulls reach sexual maturity, they separate from the family group and form loose bachelor groups for a short time, but usually stay alone. Adult bulls associate with family groups only during the mating season.
The African forest elephant forages on leaves, seeds, fruit, and tree bark of at least 96 plant species. Since it disseminates partly digested seeds for at least 5 km (3.1 mi) through its droppings, it contributes significantly to maintaining the diversity and structure of theGuinean Forests of West Africa and theCongolese rainforests.
During the 20th century, overhunting caused a sharp decline of the African forest elephant population, and by 2013 it was estimated that fewer than 30,000 individuals remained. It is threatened by habitat loss, fragmentation, andpoaching. The conservation status of populations varies across range countries. Since 2021, it has been listed asCritically Endangered on theIUCN Red List.
The African forest elephant was long considered to be asubspecies of the African elephant, together with theAfrican bush elephant. Morphological andDNA analysis showed that they are two distinctspecies.[4][5]
The taxonomic status of the African pygmy elephant (Loxodonta pumilio) was uncertain for a long time. Phylogenetic analysis of themitochondrial genome of nine specimens from museum collections indicates that it is an African forest elephant whose diminutive size or early maturity is due to environmental conditions.[6]
Phylogeny showing the relationship between living and extinct elephantids including the hybridization betweenPalaeoloxodon, represented by thestraight-tusked elephant (Palaeoloxodon antiquus) and African forest elephants
Phylogenetic analysis ofnuclear DNA of African bush and forest elephants,Asian elephants,woolly mammoths andAmerican mastodons revealed that the African forest elephant and African bush elephant are two distinct species thatgenetically diverged at least 1.9 million years ago. They are therefore considered distinct species.[7] Despite evidence of hybridization between the two species where their ranges overlap, there appears to have been littlegene flow between the two species since their initial divergence.[8]
DNA from the extinct Europeanstraight-tusked elephant (Palaeoloxodon antiquus) indicates that members of the extinct elephant genusPalaeoloxodon interbred with African forest elephants, with over 1/3 of the nuclear genome as well as the mitochondrial genome of the straight-tusked elephant deriving from that of African forest elephants, with the genomic contribution more closely related to modern West African populations of the forest elephant than to other populations.[8]Palaeoloxodon carried multiple separate mitochondrial lineages derived from forest elephants which were carried among both European straight-tusked elephant andChinese populations ofPalaeoloxodon, indicating this ancestry was widespread in EurasianPalaeoloxodon populations.[9]
Skulls of African bush elephant (left) and African forest elephant (right)
The African forest elephant is considerably smaller than the African bush elephant, though the size of the species has been subject to contradictory estimates. A 2000 study suggested that bulls of the species reach a shoulder height of 2.4–3.0 m (7 ft 10 in – 9 ft 10 in), and weighed 4–7 t (4.4–7.7 short tons), while cows were about 1.8–2.4 m (5 ft 11 in – 7 ft 10 in) tall at the shoulder and 2–4 t (2.2–4.4 short tons).[4] However, a 2003 study of forest elephants at a reserve in Gabon did not find any elephants taller than 2.16 m (7 ft 1 in).[10] A 2015 study alternately suggested that fully grown African forest elephant males in optimal condition were only on average 2.2 m (7 ft 3 in) tall and 2 t (2.2 short tons) in weight, with the largest individuals (representing less than 1 in 100,000 as a proportion of the total population) no bigger than 2.75 m (9 ft 0 in) tall and 3.5 t (3.9 short tons) in weight.[11]
The African forest elephant has grey skin, which looks yellow to reddish afterwallowing. It is sparsely covered with black coarse hair, which is 20–200 mm (0.8–8 in) long around the tip of the tail. The length of the tail varies between individuals from half the height of the rump to almost touching ground. It has five toenails on the fore foot and four on the hind foot.[12] Its back is nearly straight.[4] Its oval-shaped ears have small elliptical-shaped tips,[3] and the tip of the trunk has two finger-like processes.[13]
The African forest elephant's tusks are straight and point downwards,[4] and are present in both males and females.[13] The African forest elephant has pink tusks, which are thinner and harder than the tusks of the African bush elephant. The length and diameter vary between individuals.[12] Tusks of bulls grow throughout life, tusks of cows cease growing when they are sexually mature.[4] The tusks are used to push through the dense undergrowth of their habitat.[14] The largest tusk recorded for the species is 2.41 m (7 ft 11 in) long and 60 kg (130 lb) in weight. A larger tusk measuring 2.96 m (9 ft 9 in) long and weighing 70 kg (150 lb) has been recorded, but this may belong to a forest-bush elephant hybrid. The average tusk size is uncertain due to measurements historically being lumped in with those of African bush elephants, but based on the sizes of the largest known tusks may be in the region of 1.6–2 m (5 ft 3 in – 6 ft 7 in) and 25–30 kg (55–66 lb).[15]
A group of about 10-25 African forest elephants has been sighted on the escarpment to the east of Luanda in theKambondo forest in 2015.[24][25]
Nonetheless, it was estimated that the population of African forest elephants in Central Africa had declined by around 86% (in the 31 years preceding 2021) due to poaching and loss of habitat. In places such as Cameroon, Democratic Republic of the Congo,Republic of the Congo and theCentral African Republic, many areas of appropriate forest habitat have been reduced after years of warfare and human conflict.[20] As of 2021, an estimated 95,000 forest elephants lived in Gabon. Prior to this the population had been estimated at 50,000 to 60,000 individuals.[20]
African forest elephants in a waterholeGroup of African forest elephants digging at a mineral lickA female with her calf drinking from a spring
The African forest elephant lives in family groups. Groups observed in therain forest of Gabon'sLopé National Park between 1984 and 1991 comprised between three and eight individuals.[26] Groups of up to 20 individuals were observed in theDzanga-Sangha Complex of Protected Areas, comprising adult cows, their daughters and subadult sons. Family members look after calves together, calledallomothering. Once young bulls reachsexual maturity, they separate from the family group and form loose bachelor groups for a few days, but usually stay alone. Adult bulls associate with family groups only during the mating season. Family groups travel about 7.8 km (4.8 mi) per day and move in a home range of up to 2,000 km2 (770 sq mi).[12]Their seasonal movement is related to the availability of ripe fruits inPrimary Rainforests.[27]They use a complex network of permanent trails that pass through stands of fruit trees and connect forest clearings withmineral licks.[28] These trails are reused by humans and other animals.[29]
InOdzala-Kokoua National Park, groups were observed to frequently meet at forest clearings indicating afission–fusion society. They stayed longer when other groups were also present. Smaller groups joined large groups, and bulls joined family units.[30]
The African forest elephant feeds on bark, leaves, and fruit
The African forest elephant is anherbivore. Elephants observed in Lopé National Park fed mostly treebark andleaves, and at least 72 differentfruits.[26]To supplement their diet withminerals, they congregate at mineral-rich waterholes and mineral licks.[31][32]
Elephant dung piles collected inKahuzi-Biéga National Park contained seeds and fruit remains ofOmphalocarpum mortehanii, junglesop (Anonidium mannii),Antrocaryon nannanii,Klainedoxa gabonensis,Treculia africana,Tetrapleura tetraptera,Uapaca guineensis,Autranella congolensis,Gambeya africana andG. lacourtiana,Mammea africana,Cissus dinklagei, andGrewia midlbrandii.[33] Dung piles collected in a lowland rain forest in the northernRepublic of Congo contained seeds of at least 96 plant species, with a minimum of 30 intact seeds and up to 1102 large seeds of more than 1 cm (0.39 in) in a single pile. Based on the analysis of 855 dung piles, it has been estimated that African forest elephants disperse a daily mean of 346 large seeds per 1 km2 (0.39 sq mi) of at least 73 tree species; they transport about a third of the large seeds for more than 5 km (3.1 mi).[34]
Seeds passed by elephant gutgerminate faster. The African forest elephant is one of the most effective seed disperser in the tropics and has been referred to as the "megagardener of the forest" due to its significant role in maintaining plant diversity.[35] In theCuvette Centrale, 14 of 18 megafaunal tree species depend on seed dissemination by African forest elephants, including wild mango (Irvingia gabonensis),Parinari excelsa andTridesmostemon omphalocarpoides. These 14 species are not able to survive without elephants.[36]African forest elephants provideecological services that maintain the composition and structure of Central African forests.[37]
Vocalization is a trait found amongL. cyclotis with studies emphasizing significance in acoustic structure and their social dynamics.[38] Vocalization patterns can be classified into three main types: single rumble, single broadband, and combinatorial.[38] Rumbles are tonal, low-frequency calls, while broadband are calls that lack clear harmonic structures, resembling barks and roars.[38][39] The utilization of rumbles and broadband calls in combinatorial calls may involve distinct acoustic elements, forming multi-element calls, which combine meaningless elements to generate context-specific meaningful calls.[40][41] The African forest elephant also exhibits a more balanced distribution of combinatorial call types compared to other elephant species.[42] Despite having a simpler social structure,L. cyclotis can display a comparable repertoire of rumble-roar call combinations thanL. africana.[43] Communication patterns vary across age and sex, with adult males typically producing more combinatorial calls than adult females.[44] Additionally, certain events may provoke a behavioral change, as evidenced by lowered levels of vocalizations in response to gunfire sounds related to poaching.[45]
For these mammals, hearing and smell are the most important senses they possess because they do not have good eyesight. They can recognize and hear vibrations through the ground and can detect food sources with their sense of smell. Elephants are also an arrhythmic species, meaning they have the ability to see just as well in dim light as they can in the daylight. They are capable of doing so because the retina in their eyes adjusts nearly as quickly as light does.[46]
Females reach sexual maturity between the age of 8 and 12 years, depending on the population density and nutrition available. On average, they begin breeding at the age of 23 and give birth every 5–6 years. As a result, the birth rate is lower than the bush species, which starts breeding at age 12 and has a calf every 3–4 years.[47]
Baby elephants weigh around 105 kg (232 lb) at birth. Almost immediately, they can stand up and move around, allowing the mother to roam around and forage, which is also essential to reduce predation. The baby suckles using its mouth while its trunk is held over its head. Their tusks do not come until around 16 months and calves are not weaned until they are roughly 4 or 5 years old. By this time, their tusks are around 14 cm (5.5 in) long and begin to get in the way ofsuckling.[48]
African forest elephants have a lifespan of about 60 to 70 years and mature slowly, coming to puberty in their early teens.[49] Bulls generally pass puberty within the next year or two of females. Between the ages of 15 and 25, bulls experience "musth", which is a hormonal state they experience marked by increased aggression. The male secretes fluid from the temporal gland between his ear and eye during this time. Younger bulls often experience musth for a shorter period of time, while older bulls do for a longer time. When in musth, bulls have a more erect walk with their head high and tusks inward, they may rub their heads on trees or bushes to spread the musth scent, and they may even flap their ears, accompanied by a musth rumble, so that their smell can be blown toward other elephants. Another behavior affiliated with musth is urination. Bulls allow their urine to slowly come out and spray the insides of their hind legs. All of these behaviors are to advertise to receptive females and competing bulls that they are in the musth state. Bulls only return to the herd to breed or to socialize; they do not provide parental care to their offspring but rather play a fatherly role to younger bulls to show dominance.[50]
Females arepolyestrous, meaning they are capable of conceiving multiple times a year, which is a reason why they do not appear to have a breeding season. However, there does appear to be a peak in conceptions during the two rainy seasons of the year. Generally, the female conceives after two or three matings. Though the female has plenty of room in her uterus for twins, twins are rarely conceived. Gestation lasts 22 months. Based on the maturity, fertility, and gestation rates, African forest elephants have the capacity to increase their population by 5% annually under ideal conditions.[51]
African forest elephants are hunted by varioushunter-gatherer groups in the Congo basin, including byMbuti pygmies, among others. It is unknown how long the active hunting of elephants in the region has been practised, and it may have only begun as a response for the demand for ivory beginning in the 19th century or earlier.[52]
Elephants are traditionally hunted usingspears, typically to stab at the lower abdomen (as is done among the Mbuti) or knees, both of which are effective at rendering the animal immobile. Anthropologist Mitsuo Ichikawa observed the hunting of elephants by Mbuti pygmies in fieldwork during the 1970s and 1980s, when the Mbuti used spears tipped with metal points (though earlier reports suggest that that prior to this they used purely wooden spears, which may have been less effective at breaking the elephants' hide). As observed by Ichikawa, elephant hunting by the Mbuti pygmies involved both small and large groups of hunters, which was led by at least one experienced hunter called amtuma. Before the hunt began, ritual acts of singing and dancing were performed by the community to support the success of the hunt. These hunters often went into the forest without food, living off of wild honey and vegetables, smearing themselves in mud, elephant dung, and charcoal made from certain plants to disguise their scent from the elephants. Once the traces of an elephant are detected, it was carefully tracked, before being approached fromdownwind and stabbed. It typically took several hours to several days from the first stab to the death of the elephant.[52]
Many hunts failed due to elephants detecting the hunters before being stabbed and fleeing, with field research by Ichikawa finding that only one out of six Mbuti elephant hunts were successful in a six-month period, corresponding to around 60–70 days of total hunting time, meaning that despite the large quantity of meat provided by each individual elephant, it did not provide reliable subsistence, with the Mbuti instead relying on hunting smaller animals. Following the death of the animal, the Mbuti hunters returned to their homes, with the whole community moving to dismember the elephant carcass. Meat was shared equally among the community with the exception of a few body parts which were reserved for certain community members, with the feast on the animals remains lasting for several days. Elephant hunting was a dangerous activity that was known to result in the deaths of hunters.[52]
Both African elephant species are threatened foremost byhabitat loss andhabitat fragmentation following conversion of forests for plantations of non-timber crops,livestock farming, and building urban and industrial areas. As a result, human-elephant conflict has increased.Poaching forivory andbushmeat is a significant threat in Central Africa.[2] Because of a spike in poaching, the African forest elephant was declaredCritically Endangered by theIUCN in 2021 after it was found that the population had decreased by more than 80% over 3 generations.
Civil unrest, human encroachment, and habit fragmentation leaves some elephants confined to small patches of forest without sufficient food. In January 2014,International Fund for Animal Welfare undertook a relocation project at the request of theIvory Coast government, moving four elephants fromDaloa toAssagny National Park.[53]
Tridom is a hotspot for poaching of African forest elephants
Genetic analysis of confiscated ivory showed that 328 tusks of African forest elephants seized in thePhilippines between 1996 and 2005 originated in the eastern Democratic Republic of the Congo; 2,871 tusks seized inHong Kong between 2006 and 2013 originated in Tridom, the tri-nationalDja-Odzala-Minkébé protected area complex and the adjacent Dzanga Sangha Reserve in the Central African Republic. So did partly worked ivory confiscated between 2013 and 2014 at warehouses in Togo comprising 4,555 kg (5.021 short tons) of tusks.[54]The hard ivory of the African forest elephant makes for more enhanced carving and fetches a higher price on theblack market. This preference is evident in Japan, where hard ivory has nearly monopolized the trade for some time. Premium qualitybachi, a traditional Japanese plucking tool used forstring instruments, is contrived exclusively from African forest elephant tusks. In the impenetrable and often trackless expanses of the rain forests of the Congo Basin, poaching is extremely difficult to detect and track. Levels of off-take, for the most part, are estimated from ivory seizures. The scarcely populated and unprotected forests in Central Africa are most likely becoming increasingly alluring to organized poacher gangs.[55]
Late in the 20th century, conservation workers established a DNA identification system to trace the origin of poached ivory. Due to poaching to meet high demand for ivory, the African forest elephant population approached critical levels in the 1990s and early 2000s.[56][57] Over several decades, numbers are estimated to have fallen from approximately 700,000 to less than 100,000, with about half of the remaining population in Gabon.[58] In May 2013,Sudanese poachers killed 26 elephants in the Central African Republic'sDzanga Bai World Heritage Site.[59][60] Communications equipment, video cameras, and additional training of park guards were provided following the massacre to improve protection of the site.[61] From mid-April to mid-June 2014, poachers killed 68 elephants inGaramba National Park, including young ones without tusks.[62]
At the request of PresidentAli Bongo Ondimba, twelve British soldiers traveled to Gabon in 2015 to assist in training park rangers following the poaching of many elephants inMinkebe National Park.[63]
On 19 August 2020,Guyvanho, a poacher who killed over 500 African forest elephants in theNouabalé-Ndoki National Park, was convicted to 30 years in prison for charges of poaching and others. Guyvanho was the first poacher to be tried criminally in the Republic of the Congo, and has the longest prison sentence for a poacher in the Republic of the Congo.[64][65]
It is not ivory alone that drives African forest elephant poaching. Killing for bushmeat in Central Africa has evolved into an international business in recent decades with markets reaching New York and other major cities of the United States, and the industry is still on the rise. This illegal market poses the greatest threat not only to forest elephants where hunters can target elephants of all ages, including calves, but to all of the larger species in the forests. There are actions that can be taken to lower the incentive for supplying to the bushmeat market. Regional markets, and international trade, require the transporting of extensive amounts of animal meat which, in turn, requires the utilisation of vehicles. Having checkpoints on major roads and railroads can potentially help disrupt commercial networks.[55] In 2006, it was estimated that 410 African forest elephants are killed yearly in theCross-Sanaga-Bioko coastal forests.[66]
In 1986, the African Elephant Database was initiated with the aim to monitor the status of African elephant populations. This database includes results from aerial surveys, dung counts, interviews with local people, and data on poaching.[67]
Both African elephant species have been listed by theConvention on International Trade in Endangered Species of Wild Fauna and Flora onCITES Appendix I since 1989. This listing banned commercial international trade of wild African elephants and their parts and derivatives by countries that signed the CITES agreement. Populations of Botswana, Namibia and Zimbabwe were listed in CITES Appendix II in 1997 as was the population of South Africa in 2000.[2] Hunting elephants is banned in the Central African Republic, Democratic Republic of Congo, Gabon, Côte d'Ivoire, and Senegal.[68]
African forest elephants are estimated to constitute up to one-third of the continent's elephant population but have been poorly studied because of the difficulty in observing them through the dense vegetation that makes up their habitat.[69] Thermal imaging has facilitated observation of the species, leading to more information on their ecology, numbers, and behavior, including their interactions with elephants and other species. Scientists have learned more about how the elephants, who have poor night vision, negotiate their environment using only their hearing and olfactory senses. They also appeared to be much more active sexually during the night compared to the day, which was unexpected.[51]
Research in the tropical rainforest has shown that African forest elephants can significantly increase the forest's carbon uptake, making conservation a way to contribute tocarbon storage.[70]
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