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)

Examples of local names (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.