SPECIES VARIATION - Editorial Comment

• A variety of sub-specific designations for African elephants have been suggested, most commonly based solely on local variations in ear shape. The only real distinction has been between the savanna elephant Loxodonta africana africana and the forest elephant Loxodonta africana cyclotis. Recent genetic work has distinguished clearly between Forest and Savannah elephants, along the lines suggested by morphometric differences, suggesting that these should be considered as separate species: Loxodonta africana - African Elephant and Loxodonta cyclotis - Forest Elephant . 
  • For the major visible distinctions between these two types of elephant, see Loxodonta africana - African Elephant (Species) - General appearance - Similar Species and Loxodonta cyclotis - Forest Elephant (Species) - General appearance - Similar Species.
• Further work including analysis of elephants from West Africa suggests that the genetic relationships within Loxodonta may be complex.
• Various races or sub-species of savannah elephants named previously, mainly based only on local variations in ear shape, are not generally considered valid.

Intraspecific Variation

• DNA sequence analysis of samples from 195 free-ranging African elephants from 21 distinct populations showed that there were phylogenetic distinctions between savanna and forest elephant populations, but there were no distinct separations between different forest elephant populations. (J22.293.w1)
  • There was some evidence of limited ancestral hybridisation between Loxodonta africana - African Elephant and Loxodonta cyclotis - Forest Elephant in one population (the Garamba population) of elephants. (J22.293.w1)
• Loxodonta cyclotis - Forest Elephant and Loxodonta africana - African Elephant are distinct species, separated by a hybrid zone. (J404.37.w1)
  • A study based on three biparentally inherited X-linked gene introns, maternally transmitted mitochondrial DNA (mtDNA) and a paternally inherited Y-chromosome gene intron, found that the haplotypes for three biparentally inherited X-linked genes showed almost complete differentiation between Loxodonta cyclotis - Forest Elephant and Loxodonta africana - African Elephant. Two distinctive Y-chromosome lineages were detected, with all Forest elephant males in one clade, and all but one savanna elephant male carrying the other clade; the sole exception was from near a forest/savanna habitat transition zone. For mtDNA, again two clades were distinguished, with all in clade I being savanna elephants while those in clade II included all forest elephants plus savanna elephants from Cameroon, Tanzania, Botswana and Zimbabwe. Caution was suggested when using solely mtDNA data to determine species distinctions. (J404.37.w1)
• There are marked differences in skull and mandible morphology between the two species, with Loxodonta africana - African Elephant having larger, broader, more pneumatized crania, particularly the forehead, while Loxodonta cyclotis - Forest Elephant is wider across the skull roof and have a longer "spout" or chin region of the lower jaw. (J405.2.w1)
• Differences in appearance include: larger, more triangular ears in Loxodonta africana - African Elephant, these being smaller and more rounded in Loxodonta cyclotis - Forest Elephant; tusks which are sturdy and curve forwards and outwards in Loxodonta africana - African Elephant compared with thinner tusks directed more downwards in Loxodonta cyclotis - Forest Elephant. Loxodonta cyclotis - Forest Elephant has a more compact general body build, forequarters lower than the hindquarters, and a more vertical drop of the cranium walls behind the eyes, with a sharper backward slope of the forehead, compared with the flaring out of the cranium behind the eyes in Loxodonta africana - African Elephant. (J405.2.w1)
• There are distinct differences in skull measurements between Loxodonta cyclotis - Forest Elephant and Loxodonta africana - African Elephant, indicative of their separate status as distinct species. (J405.2.w2)
  • Loxodonta africana - African Elephant has a long skull with a high occiput; (J405.2.w2)
  • Loxodonta cyclotis - Forest Elephant has a wider postorbital constriction and a relatively broader rostrum, also a longer "spout" length: the antero-posterior length of the mandibular symphysis; (J405.2.w2)
  • Many skulls from the Parc National des Virunga, along the border between DRC and Rwanda and Uganda, and skulls from Western Uganda (Budongo Forest and West Nile District) were noted to show characteristic indicative of hybrids. (J405.2.w2)
  • Skulls from the Park National de la Garamba tended to be of one type or the other rather than showing hybrid features. (J405.2.w2)
• Recent genetic studies have supported the existence of two species of Loxodonta: Loxodonta africana - African Elephant and Loxodonta cyclotis - Forest Elephant. In a new study, using mitochondrial DNA and nuclear microsatellite markers, a more complex pattern was detected, with five clades of elephants (East and Central African savanna, West, Central and South African savannah, two West African clades and a clade containing the Central forest elephants plus two savannah elephants from Central African sites), according to phylogenetic analysis of mitochondrial DNA, while analysis of microsatellite markers found two groups of savanna elephants and two groups of forest elephants (from West Africa and Central Africa). Overall, it was suggested that there are three broad groups: forest elephants from Central African, forest and savannah elephants of West Africa, and savannah elephants of Central, eastern and southern Africa. (J179.269.w1)
• Some authorities have suggested that there are six subspecies, including four variations within the savannah elephant Loxodonta africana africana and two within the forest elephant Loxodonta africana cyclotis. (B147)
• In East Africa, three different races of elephant have been described, Loxodonta africana knochenhaueri Matschie, Loxodonta africana peeli Lydekker and Loxodonta africana cavendishi (also named by Lydekker), distinguished on the basis of ear shape. (B385.2.w2)
• As many as 17 races or subspecies have been described, including eight described in one paper by Lydekker. The following subspecies are suggested: (B384.2.w2)
  • Loxodonta africana africana (Blumenbach, 1797) the bush elephant (to include specimens formerly designated as Loxodonta africana angolensis, Loxodonta africana capensis, Loxodonta africana mocambicus, Loxodonta africana selousi, Loxodonta africana toxotis and Loxodonta africana zukowski) in South Africa, Namibia, Botswana, Mozambique, Zimbabwe, Angola, Zambia and southern Zaire. (B384.2.w2)
  • Loxodonta africana knochenhaueri (Matschie, 1900), the Maasai elephant (to include Lydekker's designated Loxodonta africana cavendishi and Loxodonta africana peeli), in Tanzania, Kenya, southwestern Somalia and most of Uganda. (B384.2.w2)
  • Loxodonta africana orleansi (Lydekker, 1907), consisting only of a small remnant in western Ethiopia (also formerly in northern Somalia). (B384.2.w2)
  • Loxodonta africana oxyotis (Matschie, 1900) "pointed ears", the savannah elephant (including Loxodonta africana rothschildi), across Africa north of the tropical forest belt, found in southern and western Ethiopia, the Sudan, the Central African republic, northern Cameroon, Chad, northern Nigeria and through Burkina Faso to northern Sierra. Loxodonta africana Leone. (B384.2.w2)