African Elephants (Loxodonta africana and L. cyclotis) Fact Sheet: Taxonomy & History

Taxonomy

• Two species of “African elephant” (Rohland et al. 2010; Shetty and Vidya 2011; Maisels et al. 2013)
  • Loxodonta africana – African savanna or African bush elephant
  • Loxodonta cyclotis – African forest elephant
  • No subspecies currently recognized (Wittemyer 2011)
    • Regional variation in body size, appearance, and ivory
• Taxonomic history and controversy
  • Formerly described as two subspecies of Loxodonta africana (Maisels et al. 2013)
  • Taxonomic designations have been much debated (Shetty and Vidya 2011)
    • See Phylogenetic Relationships

Nomenclature

• Elephant (English)
  • A possible origin (Shoshani and Shoshani 2000)
    • Means “huge arch”
    • ele, from Greek for “arch”
    • phant, from Latin for “huge”
  • Another possible origin
    • Elephant derived from the Latin elephantus and from Greek elephant- or elephas meaning "elephant, ivory" (perhaps of Hamitic origin) (Gove 1971)
• Order Proboscidea (Shoshani and Shoshani 2000)
  • From proboscis, referring to the elephant's prominent trunk
  • Pro, meaning “before” (Greek)
  • –boscis, meaning “mouth” (Greek)
  • –idea/-oidea, meaning “appearance or “kind”
  • Given by the naturalist Carl D. Illiger in the early 19^th century
• Genus: Loxodonta (Tassy and Shoshani 2013)
  • Refers to “lozenge shape of the enamel loops on the chewing surfaces of the teeth” (Shoshani 2000)
    • Asian elephant has narrow loops on its teeth
  • In 1827, an anonymous author Latinizes F. Cuvier’s French name ‘Loxodonte’ (1825)
    • Assumedly to make the genus name valid under the International Code of Zoological Nomenclature

Synonyms

• Elephas africanus [Blumencach 1797] (Wittemyer 2011)
• African elephant sometimes confused with the Asian Elephant (Elephas maximus)
  • Lives in southeast Asia

Common names

• African elephant, African savanna elephant, African bush elephant, African forest elephant (English)
• Éléphant de savane, Éléphant de forêt (French)
• Afrikanischer savannenelefant, Afrikanischer waldelefant (German)
• Elefante de sabana, Elefante de bosque (Spanish)

Other vernacular names referring to elephants (Shoshani and Shoshani 2000)

• Tembo or ndovu (Swahili)
• Benionclet (local tribes of Mount Elgon; e.g., the Elkony)
• Hastin (Sanskrit for “having a hand”)

Early elephant-like animals

• Order Proboscidea
  • About 10 living and extinct families known
  • Modern elephants belong to the suborder Elephantiformes
    • Includes elephant-like proboscideans most closely related to modern elephants
    • Known from about 34 mya
    • Characterized by specialized traits, such as a well-developed trunk and tusks
    • Early proboscideans were smaller and did not have some traits of modern elephants (eg, skeleton not adapted to carry huge weight, different tooth characteristics)
  • Rapid species diversification started around 25 to 20 mya (late Oligocene/early Miocene), followed by numerous extinctions 2 to 3 mya (Cantalapiedra et al. 2021)
  • Some proboscideans evolved to be large species, with some exceptions, such as dwarf, dog-sized island species (e.g., Elephas falconeri)
• See Evolutionary History, below

Origins of living elephants

• Divergence among living (extant) elephant species
  • Modern African (Loxodonta) and Asian (Elephas) elephants both originated in Africa approximately 6 mya (Sooriyabandara et al. 2023)
  • Diverged approximately 6 to 9 mya (Rholand 2007; Palkopoulou et al. 2018)
    • Loxodonta migrated throughout most of Africa, but not outside of Africa
    • Elephas widely dispersed from Africa to Eurasia and Asia
• Divergence of African savanna ( africana) and African forest (L. cyclotis) elephants
  • Split from each other 2 to 5 million years ago, possibly earlier (Rholand 2010)
  • Loxodonta atlantica, a larger species than africana, from the Pliocene was a widespread species that lived in northern Africa, as well as eastern and southern Africa (Tassy and Shoshani 2013)

Closest living relatives (extant species)

• Dugongs and manatees (“sea cows”) (Order Sirenia) and hyraxes (Order Hyracoidea)
  • Part of Paenungulata within Afrotheria (meaning “African mammal”) (eg, Springer 2022)

Fossil diversity

• Early fossils
  • First fossil relatives of elephants discovered in Africa, Morocco (from Gheerbrant 1996, 2009)
    • 60-million-year-old Eritherium
      • Oldest-known proboscidean
      • Weighed about 4 to 5 kg (8.8 to 11 lb)
      • Suggests an ancient origin of proboscideans and a major evolutionary leap during the Eocene (Emmanuel Gheerbrant, personal communication, 2025)
        • Larger body size, changing molar teeth shape/structure
    • 55-million-year-old Phosphatherium
      • One of the oldest proboscideans
      • More advanced than Eritherium (particularly tooth characteristics
      • Weighed about 15 kg (33 lb)
• Extinct modern elephant relatives
  • All Loxodonta fossils discovered in Africa (Tassy and Shoshani 2013)
    • Earliest known from 5 to 7 mya (Tassy and Shoshani 2013)
    • Shows early characteristic tooth enamel loops that distinguish African elephants
  • Mammoths (Mammuthus)
    • More closely related to Asian elephants than African elephants (Sukumar 2006; Palkopoulou et al. 2018)
    • Migrated out of Africa in the Late Pliocene
    • Rapidly dispersed throughout Europe, northern Asia and North America
      • Also inhabited Central America, and the British Isles and Mediterranean islands
• Mastodons
  • Diverged from other elephantids 20 to 30 mya (Shoshani et al. 2006)
  • Distantly related to modern elephants
  • Occurred in Europe, Asia, North America, and Central America
• Extinct pygmy elephants
  • Present on numerous Mediterranean islands until about 10, 000 years ago (e.g., Athanassiou et al. 2019)
  • Extinct pygmy proboscideans also known from Indonesia (e.g., Hooijer 1955), and the Philippines and Japan (Saegusa 2008; Liscaljet 2012)

Social evolution and cognition

• Elephants have large, complex brains (Roca and O’Brien 2005)
  • Share advanced traits that independently evolved across 3 mammalian orders:
    • Proboscideans (elephants, and extinct mammoths and mastodons)
    • Primates
    • Cetaceans (whales, dolphins, and porpoises)
• Social behaviors likely helped drive the evolution of their brains, and their complex thinking and sensory abilities
• See Garstang (2015) and review article by Plotnik and Jacobson (2022) for in-depth information

Books

• African elephants in culture, history, mythology, and art
  • Terrible Beauty: Elephant–Human–Ivory — U. Chicago Press (2021)
  • Thirty-Three Ways of Looking at an Elephant — Dale Peterson, editor (2020)
  • Death and Compassion: The Elephant in Southern African Literature — Dan Wylie (NYU Press, 2018)
  • Elephant — Dan Wylie (Reaktion Books, 2008)
  • Elephants: A Cultural and Natural History — Karl Gröning (1999)
  • Elephant: The Animal and its Ivory in African Culture — Doran H. Ross, editor (1992)
• Memoirs
  • An Elephant in my Kitchen: What the Herd Taught Me About Love, Courage, and Survival — Françoise Malby-Anthony (with Katja Willemsen) (2019)
  • Elephant Don: The Politics of a Pachyderm Posse — Caitlin O'Connell (2015)
  • Love, Life, and Elephants: An African Love Story — Daphne Sheldrick (2013)
  • The Elephant Whisperer: My Life with the Herd in the African Wild — Lawrence Anthony (with Graham Spence) (2009)
  • Elephant Memories: Thirteen Years in the Life of an Elephant Family — Cynthia Moss (2000)
  • Coming of Age with Elephants: A Memoir — Joyce Poole (1996)

Documentaries

• A Life Among Elephants — 2024 (Maramedia)
  • Featuring career of elephant biologist and conservationist Iain Douglas-Hamilton
• Secrets of the Elephants — 2023 (National Geographic)
  • Includes biology of African forest elephant
• The Elephant Queen — 2018 (Deeble&Stone, Apple TV)
• Mind of a Giant — 2016
• The Ivory Game — 2016 (Netflix)
• An Apology to Elephants — 2013 (HBO)
• Echo: An Elephant to Remember — 2010 (PBS)
  • Profile of an elephant named Echo and the people who cared for and studied her life
• Battle for the Elephants — 2013 (National Geographic)
  • Explores the value of African elephant tusks

Kingdom: Animalia

Phylum: Chordata

Class: Mammalia Linnaeus, 1758 — mammals

Order: Proboscidea Illiger, 1811 — elephants (and their extinct relatives, mammoths and mastodons)

Family: Elephantidae Gray, 1821 — elephants

Genus: Loxodonta (Cuvier, 1825, amended by Anonymous, 1827)

Species: Loxodonta africana Blumenbach, 1797 — African savannah or African bush elephant

Species: Loxodonta cyclotis (Matschie, 1900) — African forest elephant

Also see Taxonomy.

Sources: Tassy and Shoshani (2013), Gobush et al. (2021), Gobush et al. (2022)

African elephant; plate in B. Cuvier's 1827 The Animal Kingdom: Arranged in Conformity with its Organizations.

Image credit: © Biodiversity Heritage Library via Flickr. Some rights reserved.

Blanc (2008)
Cutler (1985)
Eggert et al. (2002)
Eltringham (1991)
Grubb et al. (2000)
Gheerbrant et al. (1996, 2009)
ITIS
Nikaido (2003)
Palombo and Villa (2001)
Poulakakis et al. (2006)
Roca et al. (2001)
Rohland et al (2007 2010)
Ross (1992)
Shoshani (1992, 2006)
Wilson & Reeder (1992)
Yang (1996)