Animal Diversity Web

Ge­o­graphic Range

The range of African ele­phants (Lox­odonta africana) is patchily dis­trib­uted across cen­tral and south­ern Africa in the Ethiopian Re­gion. Rem­nant pop­u­la­tions exist as far north­west as Guinea-Bis­sau and as far north­east as Ethiopia. Dis­junct pop­u­la­tions exist south­ward into north­ern South Africa, and in­clude con­tigu­ous pop­u­la­tions in Gabon, Tan­za­nia, Botswana, and Zam­bia. Once pre­sent in Bu­rundi, Gam­bia, and Mau­ri­ta­nia, they've now been ex­tir­pated from those coun­tries. These ele­phants were ex­tir­pated from Swazi­land, as well, but have been re-in­tro­duced here in re­cent years. (Blanc, 2008; Laursen and Bekoff, 1978; Thomas, et al., 2008)

• Biogeographic Regions
• ethiopian
  • native

Habi­tat

African ele­phants are found in many habi­tats of Africa such as sa­van­nas, rain forests, wood­lands, scrub forests, oc­ca­sion­ally deserts, and beaches. How­ever, due to poach­ing threats pro­tected sanc­tu­ar­ies are their main habi­tats. Within these sanc­tu­ar­ies, these ele­phants will in­habit areas that have sources of water and abun­dant veg­e­ta­tion for for­ag­ing. Across these habi­tats, the el­e­va­tion level ranges from sea level (0 me­ters) to 4000 me­ters. (Duffy, et al., 2011; Har­ris, et al., 2008; Laursen and Bekoff, 1978; Mash­in­to­nio, et al., 2014)

• Habitat Regions
• tropical
• terrestrial

• Terrestrial Biomes
• desert or dune
• savanna or grassland
• rainforest
• scrub forest

• Range elevation

  0 to 4000 m

  0.00 to 13123.36 ft

Phys­i­cal De­scrip­tion

Cur­rently, African ele­phants are the largest ter­res­trial or­gan­ism. Fe­male ele­phants can range from 2,000 to 3,500kg in mass and stand 2.2 to 2.6m at shoul­der height. Male ele­phants are larger, rang­ing from 4,500 to 6,100kg in mass and stand­ing 3.2 to 4m at shoul­der height. Dis­tin­guish­ing char­ac­ter­is­tics be­tween the sexes in­clude head shape, width of fore­head, sad­dle-back vs straight back, and tusk size. Males are broader with a curvy build. Both sexes have two thick, ivory tusks which are curved and can reach up to 350 cm in length. African ele­phants have mus­cu­lar trunks that are able to grab ob­jects and can be used for breath­ing pur­poses. They can grab ob­jects due to the unique shape at the end of the trunk. Their siz­able ears are tri­an­gu­lar and can help keep them cool in the hot, sum­mer months.

African ele­phants have creased, gray skin cov­ered with papil­lae. The thick (up to 30 mm), im­mo­bile skin cov­ers the ma­jor­ity of the body, while bumpy skin cov­ers areas that re­quire a lot of move­ment. Smooth skin can be found on del­i­cate areas of the body. Hair that varies in color, length, and thick­ness can be found along the body. Hair that grows along the back and tail is flat­tened dark hair that can grow to 0.80 m in length. Hair around the eyes is long in order to pre­vent for­eign ob­jects from en­ter­ing the eyes. Calves have dif­fer­ent hair col­or­ing and tex­ture. Their hair is softer with a lighter tone such as a red or brown.

The den­tal for­mula for African ele­phants is i 1/0, c 0/0, p 3/3, m 3/3. At birth, calves do not have tusks but in­stead have tem­po­rary pre­max­il­lary in­cisors. These in­cisors are re­placed at about a year of age with in­cisors that will ul­ti­mately form the tusks. Adult African ele­phants have lophodont den­ti­tion, with 6 mo­lars that grow and move for­ward, like on a con­veyor belt. They are worn down, lost, and re­placed through­out their lives. Once the sixth set of mo­lars is worn down, there are no ad­di­tional teeth to re­place them and the ele­phant is not able to process for­age. (Laursen and Bekoff, 1978; Nowak, 1999)

• Other Physical Features
• endothermic
• homoiothermic
• bilateral symmetry

• Sexual Dimorphism
• male larger
• ornamentation

• Range mass

  2000 to 6100 kg

  4405.29 to 13436.12 lb

• Range length

  2.2 to 4 m

  7.22 to 13.12 ft

Re­pro­duc­tion

The es­trus state is how bulls know if cows are ready to mate. This is done by a scent in fe­male urine and gen­i­tal area. Dur­ing the courtship process, bulls ap­proach fe­males and at­tempts to use their trunk to stroke her. Bulls will put up a fight in order to mate, by chas­ing the fe­males if they re­treat. When fe­males stop re­treat­ing, they will join the bulls in stroking each other with their trunks. The courtship con­tin­ues by fe­males sur­ren­der­ing their hindquar­ters to the males. Bulls then mount the fe­males to begin mat­ing. The males will thrust re­peat­edly into the fe­males for up to 2 mins. While fe­males are in an es­trus state, they may mate with sev­eral dif­fer­ent bulls, and are con­sid­ered polyg­y­nan­drous. They are co­op­er­a­tive breed­ers, in which fe­males have help rais­ing young by other mem­bers of the fam­ily or "clan." (Laursen and Bekoff, 1978; Poole, et al., 2007)

• Mating System
• polygynandrous (promiscuous)
• cooperative breeder

African ele­phants are vi­vip­a­rous an­i­mals, mean­ing that they birth their young live. They breed year-round with no sea­sonal dif­fer­ences. The num­ber of off­spring is usu­ally lim­ited to one for each birthing pe­riod but in rare cases, twins may be born. They breed once every 3-9 years, and will give birth to an av­er­age of four calves in their life­time. The ges­ta­tion pe­riod is about 22 months but is strongly in­flu­enced by en­vi­ron­men­tal fac­tors. The ges­ta­tion pe­riod may be shorter if the en­vi­ron­men­tal fac­tors are fa­vor­able for new­borns sur­vival. New­born African ele­phants will weigh be­tween 90-120 kg, with 100 kg being the av­er­age birth weight. Off­spring are com­pletely de­pen­dent on their mother's milk until they are weaned at four months, but con­tinue to oc­ca­sion­ally drink their mother's milk for up to three years. Young African ele­phants will gain their full in­de­pen­dence around eight years of age. Sex­ual ma­tu­rity oc­curs at dif­fer­ent ages for males and fe­males. Males will reach sex­ual ma­tu­rity around 20 years of age, while fe­males will reach sex­ual ma­tu­rity around 11 years of age. (Hilde­brandt, et al., 2006; Laursen and Bekoff, 1978; Poole, et al., 2007)

• Key Reproductive Features
• iteroparous
• year-round breeding
• gonochoric/gonochoristic/dioecious (sexes separate)
• sexual
• fertilization
• viviparous

• Breeding interval

  African elephants breed once every 3-9 years

• Breeding season

  African elephants breed throughout the year

• Range number of offspring

  1 to 2

• Average number of offspring

  1

• Average number of offspring

  1

  AnAge

• Average gestation period

  22 months

• Average gestation period

  670 days

  AnAge

• Average weaning age

  4 months

• Average time to independence

  8 years

• Average age at sexual or reproductive maturity (female)

  11 years

• Average age at sexual or reproductive maturity (female)
  Sex: female

  4018 days

  AnAge

• Average age at sexual or reproductive maturity (male)

  20 years

• Average age at sexual or reproductive maturity (male)
  Sex: male

  3650 days

  AnAge

In African ele­phant herds, pri­mar­ily the mother and other fe­males as­sist in tak­ing care of the young. When fe­male ele­phants give birth, they will move away from the herd in order to allow space for the off­spring. When they re­turn, all mem­bers of the herd will in­spect the new ele­phant thor­oughly. Until young ele­phants are around 4 years of age, they will closely fol­low their mother. Dur­ing this time, moth­ers will feed their young through breast milk and help them move around ob­sta­cles. Until young off­spring are in­de­pen­dent, around 8 years of age, they will de­pend on the rest of the herd to teach them how to use their trunks, for­age for food, and move around tough ob­sta­cles. Ma­ter­nal po­si­tion only af­fects the young if the mother is the leader of the herd, in which case, the young will be the next leader of the herd. (Laursen and Bekoff, 1978; Miller and An­drews, 2013)

• Parental Investment
• female parental care
• pre-hatching/birth
  • provisioning
    • female
• pre-weaning/fledging
  • provisioning
    • female
  • protecting
    • female
• pre-independence
  • provisioning
    • female
  • protecting
    • female
• post-independence association with parents
• extended period of juvenile learning
• maternal position in the dominance hierarchy affects status of young

Lifes­pan/Longevity

African ele­phants re­port­edly have been known to live up to a max­i­mum of 65 years in cap­tiv­ity. How­ever, un­pub­lished re­ports have stated that African ele­phants may live up to 80 years in cap­tiv­ity. In the wild, African ele­phants live for an av­er­age of 60-70 years. (Carey and Judge, 2000; Nowak, 1999; Weigl, 2005; Wiese and Willis, 2004)

• Range lifespan
  Status: captivity

  65 (high) years

• Average lifespan
  Status: wild

  60-70 years

• Average lifespan
  Status: wild

  60.0 years

  Max Planck Institute for Demographic Research

• Average lifespan
  Status: captivity

  80.0 years

  Max Planck Institute for Demographic Research

• Average lifespan
  Status: wild

  70.0 years

  Max Planck Institute for Demographic Research

Be­hav­ior

African ele­phants gen­er­ally are slow mov­ing crea­tures. Their reg­u­lar pace is 6 km/hour but, they can reach speeds of up to 24 km/hour when run­ning. How much they travel each day de­pends on the amount of re­sources in close prox­im­ity that they re­quire; the av­er­age walk­ing dis­tance for African ele­phants is around 10 km a day. Be­fore African ele­phant pop­u­la­tions were con­stricted to large reser­va­tions for pro­tec­tion, they mi­grated hun­dreds of kilo­me­ters sea­son­ally, from high to low al­ti­tudes and vice versa.

African ele­phants are ac­tive an­i­mals for the ma­jor­ity of time in a 24-hour pe­riod, due to the amount of food they must con­sume each day. They are dor­mant in the early morn­ing hours with an ad­di­tional sleep mid­day for a total of 4 hours of sleep every day. Dur­ing this time they scav­enge for food and groom them­selves. Groom­ing con­sist of African ele­phants using their trunks to cover them­selves with mud or water, and the process helps them main­tain hy­dra­tion.

Fe­male African ele­phants are so­cial an­i­mals that live in herds of 6 to 70 mem­bers. These herds have a ma­tri­ar­chal order, and con­sist of fe­males (cows) and their young. The alpha ele­phant in these herds tends to be the biggest and most dom­i­nant. Male African ele­phants (bulls) tend to only live within a herd if they are not old enough to go out on their own or for mat­ing pur­poses. Bulls will live a life in soli­tary or with a few other bulls. (Greco, et al., 2016; Laursen and Bekoff, 1978; Lee and Moss, 2012; Miller, et al., 2016; Thou­less, 1996)

• Key Behaviors
• terricolous
• diurnal
• nocturnal
• motile
• nomadic
• social
• colonial
• dominance hierarchies

Home Range

If enough es­sen­tial re­sources are pre­sent African ele­phants will move as lit­tle as 1.5 Km a day. When re­sources are scarce, African ele­phants may travel as much as 40 km a day. Thou­less (1996) re­ported that African ele­phants' home range can vary from 102 to 5527 km within a pe­riod of 25 months. (Miller, et al., 2016)

Com­mu­ni­ca­tion and Per­cep­tion

African ele­phants com­mu­ni­cate acousti­cally with oth­ers of their species. Many of their calls are low fre­quency calls of ca. 20Hz. They can make a va­ri­ety of calls in­clud­ing rum­ble, trum­pet, snort, roar, bark, and grunt. Soltis (2010) re­ports 3 other calls made by these ele­phants. They in­clude "rev, croak, and chuff." A trum­pet, roar, or growl could show signs of ag­gres­sion. A "soft chirp" shows sub­mis­sion or in­tim­i­da­tion. In­fant ele­phants will gur­gle dur­ing play and squeal when fright­ened. The African ele­phants can hear one of these calls from over 2km away. They will make these calls to warn or gather oth­ers in their herd or to sig­nal they are ready to mate. African ele­phants watch and lis­ten to their sur­round­ing en­vi­ron­ment for signs of some­thing amiss. They com­mu­ni­cate vi­su­ally by using their trunks or ears to sig­nal other herd mem­bers. Tac­tile com­mu­ni­ca­tion is be­tween a mother and her child or two ele­phants try­ing to mate. Forms of chem­i­cal com­mu­ni­ca­tion along with scent mark­ing among African ele­phants is done by males who are mat­ing with the fe­males in a clan. Around the clan the males will mark trees or bushes with their tusks or by se­cret­ing a sub­stance onto the bush. (Laursen and Bekoff, 1978; O’Con­nell-Rod­well, et al., 2006; Soltis, 2010)

• Communication Channels
• visual
• acoustic
• chemical

• Other Communication Modes
• scent marks

• Perception Channels
• visual
• tactile
• acoustic
• chemical

Food Habits

African ele­phants have a her­biv­o­rous diet con­sist­ing of tree fo­liage, fruits, herbs, grasses, and wood in­clud­ing roots, twigs, and bark. Their source of fiber comes from chew­ing on bark but not di­gest­ing the bark it­self. In order to ob­tain bark or roots, African ele­phants will over­turn a tree to for eas­ier ac­cess. Using their trunks as a tem­po­rary con­tainer or hold­ing, African ele­phants use their trunks to gather water and shoot it into their mouths. African ele­phants will feed con­tin­u­ously through­out the day, eat­ing op­por­tunis­ti­cally. They must con­sume around 50 gal­lons of water every­day in order to stay hy­drated.

African ele­phants are both browsers and graz­ers. De­pend­ing on the sea­son and lo­ca­tion, herds may de­pend more on one or the other feed­ing tech­nique. Typ­i­cally dur­ing the wet sea­sons, African ele­phants are more par­tic­u­lar about they con­sume. Dur­ing this sea­son, they are more likely to for­age on grasses. How­ever, dur­ing the dry sea­sons when food is less abun­dant, they will be more flex­i­ble with what they con­sume. Dur­ing both sea­sons, African ele­phants tend to pick food sources that are high in nu­tri­tional con­tent. (Archie, et al., 2006; Chafota and Owen-Smith, 2012; Co­dron, et al., 2006; Co­dron, et al., 2013; Laursen and Bekoff, 1978)

• Primary Diet
• herbivore
  • folivore
  • frugivore
  • lignivore

• Plant Foods
• leaves
• roots and tubers
• wood, bark, or stems
• fruit

Pre­da­tion

Due to their size, African ele­phants are not easy prey for many preda­tors. While lions (Pan­thera leo), wild dogs (Ly­caon pic­tus), hye­nas (Cro­cuta cro­cuta), and Nile croc­o­diles (Croc­o­dilus niloti­cus) are preda­tors of African ele­phants, the ma­jor­ity of these preda­tors prey on the young ele­phants that lag be­hind the rest of their group. These preda­tors will at­tempt to hunt ele­phants at night, due to the safety level de­creas­ing be­cause the herd can­not see well at night. How­ever, these preda­tors are not the prob­lem for the African ele­phant pop­u­la­tion. Hu­mans (Homo sapi­ens) hunt these crea­tures for their ivory tusks and leath­ery skin. (Jou­bert, 2006; Laursen and Bekoff, 1978)

• Known Predators

  • lions (Panthera leo)
  • wild dogs (Lycaon pictus)
  • hyenas (Crocuta crocuta)
  • Nile crocodiles (Crocodilus niloticus)
  • humans (Homo sapiens)

Ecosys­tem Roles

African ele­phants are thought to be a key­stone species, be­cause in small num­bers, they have last­ing im­pacts. They often are la­belled as bio­engi­neers. For ex­am­ple, their de­struc­tion or al­ter­ing of trees pos­i­tively in­flu­ences her­peto­fau­nal di­ver­sity, as they cre­ate more three-di­men­sional habi­tat di­ver­sity for these herps.

There are many par­a­sitic species that use African ele­phants as their host. There is a wide va­ri­ety to the kind of par­a­sites that in­fect African ele­phants such as flukes (Proto­fas­ci­ola ro­busta), ticks, blood suck­ing flies (An­thomyi­dae), round­worms (Strongy­loides pa­pil­lo­sus, Haemonchus con­tor­tus, Tri­chostrongy­lus col­u­bri­formis, Mur­shidia, Oe­sophagos­to­mum columbianum), lice, bot­flies (Pharyn­gob­o­lus africanus, Platy­cob­boldia lox­odon­tis, Rod­hain­o­myia roverei, Rut­te­nia lox­odon­tis, Neo­cutere­bra squamosa>>), war­ble flies (Hy­po­derma), pro­to­zoan par­a­sites (Babesia, Eime­ria bovis), and hook­worms (An­cy­lostoma duo­de­nale). African ele­phants do have a mu­tu­al­is­tic re­la­tion­ship with birds, as the birds will feed on the skin par­a­sites of the ele­phants, pro­vid­ing them a meal while rid­ding the ele­phants of some par­a­sites.

White egrets Bubul­cus ibis may have a mu­tu­al­is­tic rla­tion­ship with African ele­phants. These species often are seen to­gether, with the egret below or atop the ele­phant. The pre­sump­tion is that the birds are feed­ing on par­a­sites. (Baines, et al., 2015; Laursen and Bekoff, 1978; McLean, et al., 2012; Nasseri, et al., 2011; Zumpt and Wet­zel, 1970)

• Ecosystem Impact
• creates habitat
• keystone species

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

African ele­phants are used for a va­ri­ety of rea­sons that ben­e­fit hu­mans, often in­volv­ing the killing of these ele­phants. African ele­phants can sim­ply be used as large sport hunt­ing for tro­phies or they can be hunted for their tusks, ears, feet, and meat. Be­cause their tusks are made of ivory, they have been used for nu­mer­ous rea­sons such as bil­liard balls, dice, piano keys, and most com­monly, dec­o­ra­tive carv­ings. Ivory can be sold for a high price, al­low­ing some­one to live off the price of a few pounds. Their large ears are con­verted into leather to make purses. Their feet can be pre­served and made into fur­ni­ture. How­ever, given the con­ser­va­tion sta­tus of African ele­phants, these uses of ele­phant parts are dif­fi­cult to jus­tify. (Laursen and Bekoff, 1978; Wit­te­myer, 2011; Gao and Clark, 2014; Laursen and Bekoff, 1978; Wit­te­myer, 2011)

• Positive Impacts
• food
• body parts are source of valuable material

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

African ele­phants do not have many neg­a­tive eco­nomic im­pacts on hu­mans. African ele­phants de­stroy veg­e­ta­tion by using it as a food source or by knock­ing it down due to their mas­sive size. If ele­phants are raid­ing crops for food and peo­ple come to stop them, the ele­phants might chase down the peo­ple and kill them. (Gadd, 2005; Laursen and Bekoff, 1978; Thomas, et al., 2008)

• Negative Impacts
• injures humans

Con­ser­va­tion Sta­tus

Ac­cord­ing to the IUCN Red List, African ele­phants are listed as a "Vul­ner­a­ble" species. CITES ap­pen­dices list African ele­phants in both Ap­pen­dix I and Ap­pen­dix II. African ele­phants in Botswana, Namibia, South Africa, and Zim­babwe are listed in Ap­pen­dix II, while African ele­phants in other coun­tries are listed in Ap­pen­dix I. Ap­pen­dix I means the species is in dan­ger of ex­tinc­tion. This Ap­pen­dix pro­tects the species by mak­ing in­ter­na­tional trade il­le­gal when deal­ing with African ele­phants. Ap­pen­dix II states that the species is not en­dan­gered, but could be­come so if poach­ing is not reg­u­lated. The US Fed­eral List have listed African ele­phants as a "Threat­ened" species.

The rea­son some ele­phant pop­u­la­tions are in de­cline in cer­tain coun­tries is be­cause hunt­ing is legal in these coun­tries. How­ever, il­le­gal poach­ing that is not pros­e­cuted has the same neg­a­tive im­pact. Ele­phants are sold and bought legally or il­le­gally for their hides, fur, tusks, and meat. De­spite an in­ter­na­tional ban on the sale of ivory since 1989, trade in il­le­gal ivory dou­bled from 2007-2014. Or­ga­nized crime com­bined with cor­rupt gov­ern­ment of­fi­cials makes it in­creas­ingly dif­fi­cult to pun­ish of­fend­ers.

An­other rea­son for the de­cline of African ele­phants is that they are los­ing their habi­tats due to human de­vel­op­ment and ex­pan­sion.

In order to com­bat fur­ther de­cline of African ele­phants, some pop­u­la­tions have been moved to pro­tected areas to pre­vent poach­ing. How­ever, about 70% of the cur­rent range is un­pro­tected lands. So, threats are on-go­ing. Some man­age­ment ef­forts have been suc­cess­ful at in­creas­ing local pop­u­la­tions, so much so that con­tra­cep­tion or trap-and-re­lo­cate pro­grams had to be im­ple­mented to sus­tain the habi­tat. An­other hard­ship is that herds are treated dif­fer­ently across po­lit­i­cal bound­aries - the le­gal­ity of hunt­ing, the lo­cals' at­ti­tudes to­wards ele­phants, and the per­miss­abil­ity of or non-ac­tion to com­bat il­le­gal poach­ing all af­fect the pop­u­la­tions. Larger-scale con­ser­va­tion plans that cross coun­try bound­aries may ad­dress some of these is­sues. (Ben­nett, 2015; Blanc, 2008; Laursen and Bekoff, 1978)

• IUCN Red List

  Vulnerable
  More information

• IUCN Red List

  Vulnerable
  More information

• US Federal List

  Threatened

• CITES

  Appendix I Appendix II

• State of Michigan List

  No special status

Con­trib­u­tors

Meghan Howard (au­thor), Rad­ford Uni­ver­sity, Karen Pow­ers (ed­i­tor), Rad­ford Uni­ver­sity, Alex At­wood (ed­i­tor), Rad­ford Uni­ver­sity, Marisa Dameron (ed­i­tor), Rad­ford Uni­ver­sity, Tanya Dewey (ed­i­tor), Uni­ver­sity of Michi­gan-Ann Arbor.

Ref­er­ences

Archie, E., T. Mor­ri­son, C. Foley, C. Moss, S. Al­berts. 2006. Dom­i­nance rank re­la­tion­ships among wild fe­male African ele­phants, Lox­odonta africana. An­i­mal Be­hav­iour, 71/1: 117-127.

Baines, L., E. Mor­gan, M. Ofthile, K. Evans. 2015. Oc­cur­rence and sea­son­al­ity of in­ter­nal par­a­site in­fec­tion in ele­phants, Lox­odonta africana, in the Oka­vango Delta, Botswana. In­ter­na­tional Jour­nal for Par­a­sitol­ogy, 4/1: 43-48.

Ben­nett, E. 2015. Legal ivory trade in a cor­rupt world and its im­pact on African ele­phant pop­u­la­tions. Con­ser­va­tion Bi­ol­ogy, 29/1: 953-956.

Blanc, J. 2008. "Lox­odonta africana" (On-line). The IUCN Red List of Threat­ened Species 2008: e.T12392A3339343. Ac­cessed Sep­tem­ber 15, 2016 at http://​dx.​doi.​org/​10.​2305/​IUCN.​UK.​2008.​RLTS.​T12392A3339343.​en.

Carey, J., D. Judge. 2000. Longevity Records: Life Spans of Mam­mals, Birds, Am­phib­ians, Rep­tiles, and Fish. Den­mark: Odense Uni­ver­sity Press.

Chafota, J., N. Owen-Smith. 2012. Se­lec­tive feed­ing by a mega­her­bi­vore, the African ele­phant (Lox­odonta africana). Jour­nal of Mam­mal­ogy, 93/3: 698-705.

Co­dron, J., K. Kirk­man, K. Duffy, M. Spon­heimer, J. Lee-Thorp, A. Gan­swindt, M. Clauss, D. Co­dron. 2013. Sta­ble iso­tope turnover and vari­abil­ity in tail hairs of cap­tive and free-rang­ing African ele­phants (Lox­odonta africana) re­veal di­etary niche dif­fer­ences within pop­u­la­tions. Cana­dian Jour­nal of Zo­ol­ogy, 91/3: 124-134.

Co­dron, J., J. Lee-Thorp, M. Spon­heimer, D. Co­dron, R. Grant, D. De Ruiter. 2006. Ele­phant (Lox­odonta africana) diets in Kruger Na­tional Park, South Africa: Spa­tial and land­scape dif­fer­ences. Jour­nal of Mam­mal­ogy, 8/1: 27-34.

Duffy, K., X. Dai, G. Shan­non, R. Slo­tow, B. Page. 2011. Move­ment pat­terns of African ele­phants (Lox­odonta africana) in dif­fer­ent habi­tat types. South African Jour­nal of Wildlife Re­search, 41/1: 21-28.

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