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
• Species: africana
  • ‘Africa’
• Species: cyclotis

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”)

Relationships between African elephants

• Two distinct species (Rohland et al. 2010; Shetty and Vidya 2011; Maisels et al. 2013); recognized by IUCN as separate species in 2021 (Gobush, Edwards, Balfour et al. 2021; Gobush, Edwards, Maisels, et al. 2021)
  • Morphological differences (Groves and Grubb 2000; Grubb 2000; Shetty and Vidya 2011)
  • Genetic evidence (e.g., Roca et al. 2001; Roca et al. 2005; Rohland et al. 2010; Ishida et al. 2011; Brandt et al. 2012)
    • Recent studies: Forest elephants are genetically distinct (isolated)
      • Due to low female dispersal and varied reproductive success among males (Ishida et al. 2011; Brandt et al. 2012)
    • Previous confusion: Mitochondrial DNA did not show clear species-level separation in all studies (Shetty and Vidya 2011)
      • Conclusions based on mitochondrial DNA evidence alone found to be unreliable
      • Issue addressed by Ishida et al. (2011) and Brandt et al. (2012)
        • Compared rigor of mitochondrial and nuclear DNA approaches
  • Notable ecological differences
    • L. cyclotis
      • Habitat: rainforest
      • Diet: fruit important; eat a variety of plants
      • Activities affect plant germination, forest growth, and ecosystem-wide biodiversity
    • L. africana (Wittemyer 2011)
      • Habitat
        • Most habitats in Africa
        • Most abundant in dry wood/shrublands
      • Diet: grasses, roots, bark, leaves, stalks, fruits, seeds

Elephants in the fossil record

• Oldest fossils
• First elephant-like animal fossils discovered in Morocco (from Gheerbrant 1996, 2009)
• 55 million year-old Phosphatherium
• Weighed c. 15 kg (33 lb)
• 60 million year-old Eritherium
• Weighed c. 4 to 5 kg (8.8 to 11 lb)
• Fossil diversity
• Over time, there have been 10 or 11 families in the Order Proboscidea
• Characterized by modifications of the tusks, upper lips, and noses
• Evolutionary trend for size increase from dog-sized to over 4 meters (13 ft)
• Mastodons - extinct
• Group diverged from elephant ancestors 24 to 28 million years ago
• Belong to a separate family only distantly related to modern elephants
• Distribution
• Remains found in Europe, Greece, North America and Central America
• Mammoths (Mammuthus spp) - extinct
• Diversity
• Woolly mammoth (M. primigenius)
• Occupied Europe, British Isles, northern Asia, and as far south as Kansas in the United States
• After arriving in North America, some populations returned to Asia
• Columbian mammoth (M. columbi)
• Spread throughout North America into Central America
• A dwarf form of the Columbian mammoth survived on the Channel Islands of California
• Form became extinct about the time of early human contact 12,000 to 13,000 years ago
• Pygmy mammoth
• Skeletal remains found on the Mediterranean islands of Crete and Sardinia
• DNA and morphology shows closer relationship to Mammuthus than to Elephas
• Distribution
• Widespread in Europe, northern Asia, North America and central Mexico
• Not known to have occurred in South America
• Relationship to other elephant groups
• More closely related to Asian elephants than to African elephants; according to DNA studies
• Pygmy elephants
• Remains present on the islands of Cyprus and Tilos in the Mediterranean
• DNA more like modern Elephas than that of other genera
• Relationships between specimens of "pygmy elephants" on Sicily and Malta and other living and extinct elephants is unclear
• It is not known if they are more closely allied with mammoths or modern elephants

Family Elephantidae

• Estimate of family origin
  
• Originated by 16 million years ago (Mya), during the middle Miocene (Rohland et al. 2010)
• Modern Asian and African elephants likely originated in East Africa
• Living elephant diversity
• Asian elephants (Elephas)
• A new discovery of fossil remains places Asian elephants in Africa (Kenya) c. 6.7–5.2 Mya
• Descendants of the first members of this genus migrated out of Africa; north and east into Eurasia and Asia
• African elephants (Loxodonta)
• Migration from East African origins lead to populations throughout much of Africa
• Social evolution and cognition
• Elephants have large and complex brains with advanced traits that have independently evolved in only 3 mammalian orders (primates, cetaceans, proboscidea) (from Roca & O'Brien 2005)
• Social structure likely associated with increase in cognitive abilities
• All groups with complex brains  are characterized by complex social structures which require complex communication skills and advanced learning abilities

African elephants

• Estimate of origin
• Separate from Asian elephants and mammoths c. 7.6 Mya (Rholand 2007)
• Hybridization between genera can occur
• Recorded in 1979 at the Chester Zoo, England (Eltringham 1991)
• Cross between a female Asian and a male African elephant produced a calf that survived for 10 days
• Divergence of savanna and forest elephants
• Split from each other 2-5 million years ago; possibly earlier (Rholand 2010)

Closest living relatives of elephants

• Hyrax, sea cows, and golden moles
• Molecular evidence, shared anatomy, and geologic records support this close relationship
• Proposed taxa (Afrotheria) to include elephants and these animals
• Group likely originated in Africa (together with aardvarks, elephant shrews, tenrecs)

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)