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)

Rock art (from Ross 1992)

• Petroglyph images
• Representations of elephants in central Shara, Algeria, and Libya
• Dating difficult to establish, but perhaps as old as 8,000 B.C.

Ancient sculptures (from Ross 1992)

• Terracotta elephant heads
• Produced by the Nok culture of Nigeria c. 500 BCE
• The functional significance of these objects is not known

Ancient Egyptian culture (from Ross 1992)

• Hieroglyphic symbols
• Separate symbolic representations to distinguish wild elephants from trained elephants
• Used during the First Dynasty (c. 3200 BCE)
• Trade in ivory
• Egyptians participated in regional trade of ivory as early as 2420-2258 BCE

Elephants in war (from Ross 1992)

• Trained as war machines by Romans and  Egyptian pharaohs
• Hannibal's army crossed the Alps in 218-217 with elephants

Animal menageries (from Ross 1992)

• Romans displayed elephants in zoo and employed them in arena events
• Elephants served as curiosities in Egyptian, Greek and Chinese menageries

African mythology, ritual, and art

• San peoples of Western Cape, South Africa
• Gathered an elephant's power through trance
• Modern Igbo of Nigeria
• Have a long standing tradition of valuing elephants in their art and culture
• 10th century bronze elephant pendants
• Tusks found buried with prominent leaders
• Hair of elephants used for traditional anklets, necklaces
• A hat with an attached elephant tail signifies an important person to Lega of Zaire
• Elephant masks
• Traditions of wearing elephant masks found in at least 40 African peoples
• Used for sacred and secular purposes

Exploitation by humans

• Not restricted to more modern times
• By 7th century elephants became locally extinct in northern Africa (Cutler 1985)
• Driven by demand for elephant ivory and for their use as machines of war and performing animals in public spectacles

Popular culture resources

• Documentaries
• Battle for the Elephants - 2013, National Geographic Film
• The film explores the value of African Elephants tusks
• Elephant Diaries - 2005 and 2007, BBC and Animal Planet; each series contains 5 half hour episodes
• Series 1 - A year in the life of a group of orphaned baby elephants recently rescued in Kenya
• Series 2 - A year in the life of a unique group of orphaned African Elephants in Kenya
• Echo: An Elephant to Remember - 2010, PBS
• This hour long episode from season 29 of Nature takes a look at Echo's extraordinary life through footage and interviews with researchers that cared for and studied her
• Echo and Other Elephants - 2008, BBC; 8 television programs on 2 discs
• David Attenborough narrates and chronicles the work of elephant expert Cynthia Moss in Kenya's Amboseli National Park
• An Apology to Elephants - 2013, HBO
• Documentary that explores the abuse of these animals and shows how some people are reversing the tren

• Other appearances
• Mpala Live! short clip of an African Elephant foraging

• Research Memoirs
• Coming of Age with Elephants: A Memoir - Joyce Poole, 1996
  
• Animal behaviorist Poole includes her work with 800 elephants at Kenya's Amboseli National Park and all her discoveries.
• Elephant Memories: Thirteen Years in the Life of an Elephant Family - Cynthia Moss, 2000
• Also taking place at Kenya's Amboseli National Park, Moss' long-term research has revealed a lot as this book chronicles the lives of many elephant familes throughout her 27 years at the Park.
• Love, Life, and Elephants: An African Love Story - Daphne Sheldrick, 2013
• This memoir takes a look at how Daphne has saved countless African animals, including saving many newborn African Elephants from certain death.
• Elephant Don: The Politics of a Pachyderm Posse - Caitlin O'Connell, 2015
• A rare inside look at the social world of African male elephants is observed.
• Elephant Sense and Sensibility - Michael Garstang, 2015
• A comprehensive treatment of the full range of elephant behavior, including anecdotal and photographic material.

Kingdom: Animalia

Phylum: Chordata

Class: Mammalia

Order: Proboscidea

Family: Elephantidae

Subtribe: Loxodontina

Genus: Loxodonta

Species: Loxodonta africana (savannah or African bush elephant)
Species: Loxodonta cyclotis (African forest elephant)

Describer: Order established = Illiger (1811). Family describer = Gray (1821). Revised by Maglio (1973). The Asian elephant (Elephas) = Linnaeus (1758). African elephant (Loxodonta) = Cuvier (1825) L. africana = Blumenbach, (1797) and L. cyclotis = Matschie (1900)

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)