African elephant

African elephants Temporal range: Late Miocene – Present, 7–0 Ma PreꞒ Ꞓ O S D C P T J K Pg N
African bush elephant bull in Kruger National Park
African forest elephant in Nouabalé-Ndoki National Park
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Proboscidea
Family:	Elephantidae
Tribe:	Loxodontini
Genus:	Loxodonta Anonymous, 1827[1]
Type species
Elephas africana[1] Blumenbach, 1797
Species and subspecies
L. africana L. cyclotis † L. adaurora † L. a. adaurora † L. a. kararae † L. atlantica † L. a. angammensis † L. a. atlantica † L. exoptata † L. cookei
Range of living Loxodonta (2007)

African elephants comprise two extant elephant species in the genus Loxodonta: the larger African bush elephant (L. africana) and the smaller African forest elephant (L. cyclotis). Both are highly social, grey-skinned herbivores, yet they can be distinguished by body size, ear and skull shape, and the size, curvature and colour of their tusks.

As of 2021 the IUCN Red List classifies the bush elephant as Endangered and the forest elephant as Critically Endangered, citing accelerating habitat loss and fragmentation together with intense poaching for the illegal ivory trade in many range states.^[2][3]

Loxodonta is one of two surviving genera in the family Elephantidae. The generic name derives from the Greek words λοξός (loxós, “slanting”) and ὀδούς (odoús, “tooth”), referring to the lozenge-shaped enamel ridges on the molar teeth.^[1] Fossil remains attributable to Loxodonta have been documented across Africa from the Late Miocene—about 7 million years ago—to the present, tracing a once-diverse lineage that culminated in today’s two species.^[4]

The generic name Loxodonta was coined by French zoologist Georges Cuvier in 1825.^[5] It combines the Ancient Greek words λοξός (loxós, "slanting, oblique") and ὀδούς (odoús, "tooth"), a reference to the lozenge-shaped enamel plates on African elephant molars. These diamond-shaped ridges differ markedly from the more rounded enamel lamellae of the Asian elephant genus Elephas.^[4]

The African elephant first entered Western zoological literature in 1797, when German naturalist Johann Friedrich Blumenbach described it as Elephas africanus in the fifth edition of his Handbuch der Naturgeschichte.^[6]

The generic name Loxodonte was introduced by French zoologist Frédéric Cuvier in 1825 for the African elephant, drawing attention to its distinctive lozenge-shaped molar plates. Two years later an anonymous reviewer Latinised the name to Loxodonta, a spelling that has since gained universal acceptance.^[7] The International Code of Zoological Nomenclature formally confirmed the anonymous 1827 author as the nomenclatural authority for Loxodonta in Opinion 1926 (1999).^[1]

Elephas (Loxodonta) cyclotis was described by German zoologist Paul Matschie in 1900 from three Cameroonian specimens whose skulls were shorter, broader and more concave-fronted than those of other African elephants.^[8] Glover Morrill Allen treated Matschie’s taxon as a distinct species—the “forest elephant’’—in 1936,^[9] but most later authors relegated it to subspecific status within the bush elephant.^[10][11]

Consensus shifted again in the early 2000s, when comparative morphology and multilocus genetics showed deep evolutionary separation between the savanna-dwelling bush elephant and the rainforest-dwelling forest elephant, supporting full species rank for each.^{[12][13][14][15]}

Richard Lydekker had earlier proposed six ear-based subspecies in 1907,^[16] but these taxa are now regarded as synonymous with the African bush elephant.^[1]

A third putative species—the so-called West African elephant—remains unconfirmed. Mitochondrial and microsatellite data suggest that West African populations have been isolated for roughly 2.4 million years, yet additional genomic and morphological evidence is required before taxonomic recognition.^[17]

Several extinct Loxodonta taxa have been described from African fossil sites dating from the Late Miocene through the Pleistocene:

• North African elephant († Loxodonta africana pharaohensis) — named by Paulus Edward Pieris Deraniyagala in 1948 from a specimen recovered in the Faiyum Depression, Egypt.^[18]

• † Loxodonta atlantica — originally published as Elephas atlanticus by Auguste Pomel in 1879 on the basis of a skull and associated bones from Ternifine, Algeria.^[19]

• † Loxodonta exoptata — erected by Wilhelm Otto Dietrich in 1941 for molar teeth unearthed at Laetoli, Tanzania.^[20]

• † Loxodonta adaurora — a near-complete skeleton from Kanapoi, Kenya, described by Vincent Maglio in 1970.^[21]

• † Loxodonta cookei — established by William J. Sanders in 2007 from dental remains in the Varswater Formation at Langebaanweg, South Africa.^[22]

Relationships among living and extinct elephants inferred from whole-genome data are summarised in the cladogram below, after Palkopoulou et al. 2018.^[23]

The earliest named species in the genus Loxodonta is †Loxodonta cookei, known from dental and skeletal material dated to about 7–5 Ma in Chad, Kenya, Uganda and South Africa.^[24]

Analyses of nuclear DNA sequences indicate that the two extant African elephant lineages diverged c. 2.6–5.6 Ma, with the forest elephant retaining particularly high genetic diversity—probably the legacy of repeated rainforest contractions and expansions during the Pleistocene.^[14]

Sampling from 21 localities across the savanna–forest ecotone shows limited ancient gene flow: several bush elephants possess forest-elephant mitochondrial DNA, implying historical hybridisation in contact zones, yet nuclear introgression has remained minimal since the initial split.^[25]

Whole-genome sequencing of the European †straight-tusked elephant Palaeoloxodon antiquus reveals that the genus Palaeoloxodon is more closely related to African than to Asian elephants, and that extensive hybridisation occurred with L. cyclotis: over 30 % of the nuclear genome and the entire mitochondrial genome of P. antiquus derive from the forest-elephant lineage, with an affinity strongest to modern West African populations.^[23] Subsequent work on Chinese Palaeoloxodon mitogenomes suggests this forest-elephant ancestry was widespread throughout the genus.^[26]

African elephants have grey, deeply folded skin up to 30 mm (1.2 in) thick, sparsely covered with dark-brown to black bristles. Short tactile hairs line the trunk, which ends in two finger-like projections—Asian elephants have only one.^[10]

The species’ outsized ears function as thermal radiators: flapping produces air currents and exposes the highly vascular inner surface, accelerating heat loss.^[27]

The trunk—an elongation of upper lip and nose—is powered by an estimated 40,000–60,000 muscle fascicles and innervated mainly by the trigeminal nerve. It can lift about 3 % of the elephant’s body mass and is used for olfaction, touch, feeding, drinking, dusting, vocalisation, carrying, defence and attack.^[27] Elephants are competent swimmers and hold the trunk above water as a snorkel.^[28][29]

Permanent tusks—elongated upper incisors—begin to emerge when calves are roughly a year old, replacing the small deciduous “tushes’’ present at birth. Made almost entirely of dentine, tusks display diamond-shaped growth structures nearer the centre that enlarge towards the periphery.^[27] Typical tusks weigh 23 to 45 kg (51 to 99 lb) and measure 1.5 to 2.4 m (4.9 to 7.9 ft), curving gently forward as they grow throughout life. They are used for digging roots, debarking trees, sparring during musth and fending off predators.^[30]

The dental formula is 1.0.3.30.0.3.3 × 2 = 26.^[27] Each of the four functional molars weighs about 5 kg (11 lb) and is roughly 30 cm (12 in) long. As the anterior pair wears out it is shed in pieces; the posterior pair then migrates forward and a new molar erupts behind it. This horizontal tooth replacement occurs four to six times, and loss of the final set (usually at 40–60 years) often results in starvation. Compared with Elephas, Loxodonta molars have fewer enamel lamellae; their wear surfaces form the distinctive lozenge (loxodont) pattern that inspired the genus name.^[31] Some extinct Loxodonta retained permanent premolars, but these teeth are absent in the two living species.^[32]

The African bush elephant is the world's largest land animal. Adult females in prime habitat stand 2.5 to 2.7 m (8.2 to 8.9 ft) at the shoulder and weigh 2,600 to 3,500 kg (5,700 to 7,700 lb). Adult bulls average 3.0 to 3.4 m (9.8 to 11.2 ft) tall and 5,200 to 6,900 kg (11,500 to 15,200 lb) in mass, with the largest verified individual measuring 3.96 m (13.0 ft) and an estimated 10,400 kg (22,900 lb).^[33] The species’ back is concave, whereas that of the African forest elephant is nearly straight.^[12]

African forest elephants are considerably smaller; mature males attain about 2.1 to 2.3 m (6.9 to 7.5 ft) in height and weigh between 1,700 and 2,300 kg (3,700 and 5,100 lb) under favourable conditions.^[33]

African elephants occupy a patchwork of habitats across Sub-Saharan Africa. The bush elephant ranges from the Sahel southwards to northern South Africa, occurring in semiarid scrub, savanna, mopane and miombo woodlands, seasonally flooded grasslands, and montane forests up to about 4,000 m (13,000 ft) in the Rwenzori Mountains.^[34][35]

The forest elephant is restricted to the dense, evergreen and semi-deciduous rainforests of Central and West Africa, with strongholds in the Congo Basin and fragmentary populations from southeastern Nigeria to western Uganda.^[36] Both species make seasonal or altitudinal movements that track rainfall, fresh forage and, in arid zones, permanent water.

Wild African elephants are among the least-sleeping mammals known. Biologging studies on free-ranging matriarchs in the Okavango Delta recorded an average of just 120 ± 30 minutes of sleep per 24-hour cycle, usually taken standing during the predawn hours; individuals sometimes went two consecutive nights without measurable sleep.^[37] This extreme polyphasic schedule is thought to reflect predation risk, ecological pressures to feed for 16-plus hours daily, and thermoregulatory constraints in hot climates. Captive elephants, by contrast, typically sleep three to seven hours per day, much of it recumbent.^[38]

African elephants live in **matriarchal family units** composed of related adult cows, their juvenile daughters and pre-pubescent sons. The senior cow, or **matriarch**, guides the group’s movements and draws on decades-long social and ecological knowledge.^[39] Forest-elephant families are smaller and less cohesive than those of savanna elephants, probably owing to lower predation pressure and the patchy resources of dense rainforest.^[40]

Several families that share genetic ties often gather into **bond groups**, and these in turn may coalesce into larger clans at water sources or mineral licks. Males leave the natal family around puberty to form loose **bachelor associations** that establish dominance hierarchies and provide social learning.^[41]

Contrary to the long-held view that bulls live solitary lives, GPS-tracking studies show that older males act as **repositories of spatial knowledge**, leading younger companions to seasonal food and water and mentoring them through sparring and affiliative trunk-touching.^[42] Mature bulls re-enter female groups mainly for courtship and mating.

Elephants communicate through a rich repertoire of trumpets, rumbles and infrasonic calls (<20 Hz) that travel several kilometres through air and ground. Individuals recognise the voices of hundreds of conspecifics, even after years of separation, demonstrating an exceptionally wide social memory.^[43][44] Courtship often begins with soft rumbles and the gentle entwining of trunks.^[45]

African elephants are **mixed feeders** that strip foliage with the trunk and lever or pry branches loose with their tusks, often causing extensive damage to woody vegetation.^[30] Their massive hindgut provides ample fermentation space, allowing rapid throughput of bulky forage and enabling daily intakes that far exceed those of fore-gut fermenters of similar size.^[46] Thanks to this strategy they can subsist on a wide spectrum of plant parts—fibrous stems, bark, roots, grasses and fruit—selecting items according to seasonal availability.^[47]

Like their Asian relatives, **African elephants rank among the most cognitively complex non-human animals**. Their brains—averaging just over 5 kg (11 lb) and containing as many cortical neurons as a human’s—exhibit an unusually folded neocortex and enlarged temporal lobes associated with long-term memory and auditory processing.^[48][49]

• • Self-recognition and empathy.** Although the landmark mirror-test study was conducted on an Asian elephant,^[50] subsequent observations show African elephants inspect mirror marks on their own bodies and console distressed herd-mates, behaviours interpreted as self-awareness and empathy.^[51]

• • Long-distance communication and language‐like discrimination.** Field playback experiments demonstrate that savanna elephants:
• Distinguish the voices of up to 100 individual conspecifics.^[41]
• Differentiate human languages, age-classes and sex, reacting more defensively to the voices of Maasai men—traditional spear hunters—than to Kamba farmers.^[52]
• Produce a distinct low-frequency “bee alarm” rumble that prompts rapid, cohesive retreat when simulated honey-bee sounds are played back.^[53]

• • Problem solving and tool use.** Captive African elephants have succeeded in cooperative rope-pull tasks that require understanding of partner effort,^[54] manipulated water to retrieve out-of-reach food, and fashioned branches to swat flies or plug drinking holes—evidence of foresight and flexible tool use.^[55]

• • Memory and spatial mapping.** GPS studies show matriarchs recall the locations of distant waterholes and drought-refugia used decades earlier, steering their herds along optimal routes during dry spells.^[56]

Collectively, these findings place African elephants on a cognitive tier comparable with great apes and cetaceans, illustrating **convergent evolution of complex intelligence** in large-brained, socially driven mammals.^[57]

Bull elephants sparring in mock aggression

Mating ritual

African elephants reach peak reproductive output between 25 and 45 years of age. Females enter oestrus for 2–7 days at irregular intervals, with longer anoestrous phases during droughts or food shortage.^[58] A single calf is born after a gestation of about 22 months—the longest of any living mammal—and mothers typically calve once every 3–6 years, yielding up to seven offspring over a lifetime.^[59] Neonates weigh roughly 100 kg (220 lb) and are reared not only by the mother but also by related young females, a system of **allomothering** that improves calf survival.^[39]

Marked sexual dimorphism arises by the early 20s owing to faster growth in males; by 25, bulls average twice the mass of cows, yet both sexes continue to gain height and weight throughout life.

From the mid-teens, males experience annual or biennial episodes of **musth**—a testosterone-fuelled state lasting days to months and signalled by temporal-gland secretion, urine dribbling and heightened aggression.^[60] Bulls in full musth dominate mating, siring the majority of calves; outside musth they father few.^[61]

Estrous cows advertise receptivity with low-frequency calls and scent cues detectable kilometres away. They may be pursued by several males, but guard association is usually monopolised by an older, high-ranking bull in musth.^[62][60] Younger or non-musth males gain occasional matings during the early and late stages of oestrus when guarding is lax.

Mating success rises steeply from the early 30s, peaks around 40 and declines thereafter, reflecting both body size and social rank.^[63] Larger sires tend to produce heavier calves, suggesting continuing sexual selection for body mass.^[64] Bulls usually remain with a receptive herd for about a month before resuming wide-ranging searches for other fertile females, a strategy favoured by the species’ low female oestrus synchrony.

African elephants face a combination of **habitat loss and fragmentation, ivory poaching, human–elephant conflict and climate-related pressures**. The IUCN Red List (2021 –2022) classifies the savanna elephant as Endangered and the forest elephant as Critically Endangered.^[65][66]

Modelling of historical range suggests that up to 27 million elephants roamed Africa in the early 1800s; by 2024 the continental total is estimated at **≈415,000 individuals**, a loss of more than 98 %.^[67]

Poaching peaked in 2011 when the MIKE programme recorded a **Proportion of Illegally Killed Elephants (PIKE)** of 0.77. PIKE has fallen steadily, reaching its lowest level since monitoring began in the 2023 analysis (0.35 continent-wide), yet remains well above the sustainability threshold in Central and West Africa.^[68]

• **Drivers.** Rising ivory prices in the 1970s–1980s and again after the partial reopening of trade in 2006 fuelled organised poaching gangs, particularly in the Congo Basin and the Selous–Niassa corridor.^[69]
• **Policy shifts.** China, once the largest legal market, closed its domestic ivory trade in 2017; five-year reviews show a sharp drop in retail prices and a 71 % fall in seizures linked to Chinese consumers.^[70] The EU tightened import and re-export rules in 2022 and opposes any future loosening of CITES ivory controls.^[71]
• **Current hotspots.** Poaching remains acute in parts of the eastern DRC, northern Mozambique and Cameroon’s TRIDOM landscape, where weak governance and armed conflict impede enforcement.

Agricultural expansion, logging, mining and infrastructure have removed or degraded over two-thirds of elephant range since 1950. Carrying-capacity modelling suggested only 9 million elephants could be accommodated by the late 1980s;^[72] subsequent land-use change has reduced this further, especially in West Africa where fewer than 8 % of historical forest-elephant range remains intact.^[73]

Increasing frequency of extreme droughts causes die-offs and brings elephants into greater conflict with farmers. In 2024 Namibia authorised the cull of 83 elephants in over-crowded drought-stricken areas as water and forage failed.^[74]

As people settle closer to elephant habitat, crop-raiding and property damage intensify. Simple, low-cost deterrents such as beehive fences and chili-grease barriers have reduced conflict incidents by up to 80 % in field trials.^[75][76]

Poaching levels have declined since their 2011 peak, and savanna-elephant numbers are stabilising in parts of southern Africa, yet overall populations continue to fall where law enforcement is weak and land conversion rapid.^[77] Sustained anti-poaching funding, trans-boundary habitat corridors, and community-led coexistence initiatives remain critical to the long-term survival of both African elephant species.

The **African Elephant Database (AED)**, launched in 1986, remains the principal clearing-house for aerial counts, dung surveys, telemetry and poaching data across the 37 range States.^[78] A digital portal released in 2023 integrates MIKE and ETIS records and provides near-real-time dashboards for law-enforcement agencies.^[79]

Two continent-scale surveys illustrate contrasting regional trends.

• The **Great Elephant Census** (2016) counted 352 271 savanna elephants in 18 countries, documenting a 30 % decline between 2007 and 2014, with sharp losses in Mozambique and Tanzania.^[80]
• An aerial census of the **Kavango–Zambezi (KAZA) Transfrontier Conservation Area** in 2022 estimated a stable aggregate of ≈ 228 000 elephants, the world’s largest contiguous population.^[81]

African elephants were transferred to **CITES Appendix I** in 1989, halting most international ivory commerce. The listing, coupled with domestic moratoria, triggered recoveries in parts of southern Africa where protection was strong.^[82] China—formerly the largest legal ivory market—closed commercial sales in 2017, and five-year reviews show a 71 % fall in seizures linked to Chinese consumers.^[83] At **CITES CoP19** (2022) Parties reaffirmed a near-total prohibition on exporting wild-caught elephants to zoos, limiting removals to “exceptional in-situ conservation benefits”.^[84]

The **AfESG Human–Elephant Conflict Task Force** disseminates low-cost deterrents such as beehive fences and chilli-grease barriers, which field trials in Kenya and Tanzania cut crop-raiding incidents by up to 80 %.^[85][86] Community-based models, notably Namibia’s 86 conservancies, have **tripled** the national elephant population to ≈ 24 000 over two decades while generating tourism income.^[87]

Twelve West African nations adopted a **West African Elephant Memorandum of Understanding** in 2005, with CMS funding the regional strategy.^[88] Large-scale corridors such as **KAZA TFCA** and the **Great Limpopo Transfrontier Park** maintain genetic connectivity for > 300 000 elephants across five countries. A 2024 summit endorsed joint anti-poaching patrols and drought-response funds.^[89]

In 2019 CITES Parties banned commercial export of wild African elephants to zoos, except under strict, conservationally justified conditions—a measure backed by the EU but opposed by Zimbabwe and Botswana.^[90]

High elephant densities reduce woody biomass but enhance **soil carbon sequestration** through litter incorporation and nutrient cycling, offsetting above-ground carbon losses and aligning elephant conservation with climate-mitigation goals.^[91]

Many African societies revere the elephant as an emblem of **strength, longevity and wisdom**.^[92]

Myth-makers highlight its size and cooperative nature; oral traditions across West, Central and Southern Africa credit elephants with unrivalled memory and moral authority, frequently casting them as judges in animal fables.^[93] In several totemic belief systems, chiefs are thought to be **reincarnated** as elephants, reinforcing royal legitimacy and continuity.^[94]

Archaeological evidence attests to early ritual importance: 10th-century burials at Igbo-Ukwu (Nigeria) contained elaborate regalia and multiple elephant tusks, signalling high status and spiritual potency.^[95]

South Africa’s national coat of arms incorporates twin elephant tusks symbolising *wisdom, strength, moderation and eternity*.^[96]

In the 19th-century Kingdom of Dahomey, the elephant was the personal emblem of Kings Guezo and Glele, evoking power and dynastic memory; court paraphernalia and regimental banners often bore a crowned elephant motif.^[97][a]

The elephant features prominently in the heraldry of several African states:

• **Central African Republic** – elephant head in the upper-left quadrant of the shield.
• **Côte d’Ivoire** – central elephant head on the national coat-of-arms.
• **Republic of the Congo** – two elephants as supporters of the shield.
• **Guinea (1958–1984)** – golden elephant on the shield.
• **Eswatini** – elephant and lion supporters.

Collectively these emblems represent **strength, vigilance and national heritage**.^[98]

• Africa's Elephant Kingdom
• Indian elephant
• List of individual elephants
• Sri Lankan elephant
• Sumatran elephant

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