Plains Zebra (Equus quagga) Fact Sheet: Behavior & Ecology

Diurnal (from Klingel 2013 unless otherwise noted)

• Active in daylight
• Travel, feed, socialize, and rest
• Feed for long periods
  • c. 50% of time
  • > 80% of time when conditions are poor (Estes 2012)
• Visit waterholes, almost daily (Smuts 1975)
• Congregate to rest at night
• Nighttime activity is not uncommon (Estes 2012)

Daily activity patterns (from Klingel 2013 unless otherwise noted)

• Morning
• Marked by movement to grazing areas or water
• Grazing interrupted with intervals of rest
  • Up to 3 periods per day
  • Stand to nap (Estes 2012)
• Night
• Sleep up to 7 hours (Estes 2012)
  • Lie flat on the body side or semi-recumbent with legs gathered (Estes 2012; Klingel 2013)
    • Often sleep very soundly
  • 1 or more group members keeps watch; snort to rouse herd members (Estes 2012)

Annual activity patterns (from Klingel 2013 unless otherwise noted)

• Seasonal migration by some populations
  • Track vegetation surges caused by regional rainfall (Hack et al. 2002; Harris et al. 2009; Maddock 1979)
  • Migratory populations
    • Tanzania and Kenya, between the Serengeti and Masai Mara
    • Namibia and Botswana, between the Chobe and Nxai Pan
    • Botswana, across the Kalahari
    • Tanzania, across Tarangire National Park
  • Migratory distances: c.110-500 km (68-311 mi) round-trip (Harris et al. 2009; Naidoo et al. 2014)
    • Serengeti-Masai Mara ecosystem: c. 200-400 km (124-249 mi) (Harris et al. 2009)
    • Chobe-Nxai Pan ecosystem, Namibia/Botswana: c. 500 km (311 mi) or more (Naidoo et al. 2014)

Seasonal ranges (from Klingel 2013 unless otherwise noted)

• Migratory populations shift location
• Dependent on availability of food
• Serengeti-Masai Mara ecosystem, Tanzania and Kenya: 400-600 km² (249-373 mi²)

Annual ranges (from Klingel 2013 unless otherwise noted)

• Size highly variable across range
  • Kruger National Park, South Africa: 49-566 km² (19-219 mi²) (Smuts 1975)
  • Serengeti-Masai Mara ecosystem, Tanzania and Kenya: > 2000 km² (> 772 mi²)
  • Ngorongoro Crater, Tanzania: 80-250 km² (50-155 mi²)
• Focus activity near water supplies (Smuts 1975; Klingel 2013)

Highly social animals (from Klingel 2013 unless otherwise noted)

• Live in harem (family) groups
• Harem structure
• A single adult stallion (male), one or more adult mares (females), and their offspring (<3 years of age) (Hack et al. 2002; Klingel 1965)
  • Stallion leader 4.5-12 years of age (Smuts 1976c)
  • Adult membership typically stable
    • Stallion leader remains until death or unseated by younger males when old, weak, or wounded
    • Mares remain within a single group after reaching sexual maturity (Klingel 1965)
      • Rarely disperse to other groups
      • Membership not forced by stallion leader; remain together even with death of leader stallion (Klingel 1965)
• Form large herds of several hundred individuals (Klingel 1969)
  • Many harems live in close proximity to one another (Klingel 1967; Schilder 1988)
    • Individuals congregate to sleep and graze (Klingel 1969)
  • Aggregations are anonymous; harems live independently of one another
    • Subdivide and merge in a haphazard fashion throughout the year (Klingel 1969)

Harem size (from Klingel 2013 unless otherwise noted

• Regionally variable
• At most 6 adult females join a leader stallion
• Average harem sizes (including adults and juveniles): 4.2-8.2 (Hack et al. 2002; Smuts 1976c)
• Largest harem reported (indlucing adults and juveniles): 16; Ngorongoro Crater, Tanzania (Hack et al. 2002)

Dominance hierarchy linear (from Klingel 2013 unless otherwise noted)

• Leader stallion
  • Alpha individual within the harem
  • Defends group against competitors and predators
  • Sought out by lost subadult, group members
• Alpha mare
• Leads group movements
• Dominance rank determines spatial distribution of adult mares
• Foals walk behind their respective mothers
• Young male stallions
• Hold equal ranks with one another within the harem

Bachelor/stallion groups

• Young males disperse from natal groups, remain alone or join bachelor groups (Klingel 1965; Smuts 1976c)
  • Size of bachelor groups: up to 15 (Klingel 1965)
  • Smaller groups of males may remain together for extended periods, multiple years (Klingel 1965)

• Non-territorial (Klingel 2013)
• Home ranges overlap and shift seasonally (Klingel 2013)

Aggression (from Klingel 2013 unless otherwise noted)

• Threat displays 
  • Ears pricked and/or held back with head held high (Estes 2012)
    
    • Teeth often bared (Estes 2012)
  • High-step (Estes 2012)
  • Arched tail (Estes 2012)
• Fights unritualized
  • Similar to that of other equids
• Bite and chase
  • Attacks directed toward the head, neck, ears, and legs of one's opponent
  • Blunt incisors typically cause minor wounds, e.g. to the ears
• Strike out with hooves
  • Rear up on hindlegs; forelegs flail at opponent in an attempt to put him off balance
• Kick out with hindlegs in defense when fleeing
• Neck wrestle
  • Each tries to get his neck above that of his opponent
  • Goal is to unbalance one's adversary

Affiliative behaviors (from Klingel 2013 unless otherwise noted)

• Familial affection
  • Family members care for and look after one another
• Group movement often held up for the sick, weak, or injured
• Lost or lagging members may be led back to the group
• Social grooming
  • Mutually clean one another's coat and mane
  • Teeth nibble and scrape the neck, shoulders, back and sides
  • Click here for a brief video
• Close standing and head resting (Estes 2012)
  • Pair stands head-to-tail or looking over one another's shoulder
    • Tails swish flies from partner
  • Head rests on partner's back
• Greeting ceremony
  • Naso-nasal contact between non-familial stallions
    • Stallions leave their families several times per day to interact with males of nearby groups
    • Nuzzle and sniff one another
      • Necks stretched, ears directed forward, corners of mouth drawn up, and chew with bared teeth
      • Genital sniffing often incorporated between nasal contacts
  • No overt aggression apparent in such encounters
• Defecation ceremony
  • Stallions gather around the droppings of a fellow male
  • Participants sniff, lip-curl (flehmen), and defecate, producing a communal dung pile
  • Play and naso-nasal greeting ceremony often incorporated

Play (from Waring 1983 unless otherwise noted)

• Similar to domestic horses
• Infants (foals)
  • Exaggerated withdrawal and approach; galloping play
    • Circle, swerve, buck, jump, and kick
  • Nip, nibble, paw, and kick at mothers
  • Peer relationships develop as maternal focused play gives way to play with other foals or yearlings
• Locomotor play
  • Run with exaggerated movements, alone or in groups
  • Kick, jump, and buck
• Aggressive play
  • Chase and circle one another
  • Bite rival on the neck and head or legs
  • Push opponent off balance
  • Rear up on hindlegs and strike with forelegs
• Other play patterns
  • Mutual grooming; gently nibble partner with the mouth
    • Often observed between bouts of more aggressive play
  • Toss and manipulate objects with the mouth
  • Paw objects with the hooves
  • Simulated sexual mounting; more common among colts (young males) than fillies (young females)

Vocalization (from Klingel 2013 unless otherwise noted)

• Calls of alarm and greeting
  • Made by adults and foals
  • Calls of individuals are unique and distinguishable by a human observer
    • Family members likely recognize one another by sound
• Contact call
  • Primary sound produced by exhaling
  • One-to three-syllable barking sound
    • 'quaha, quaha quaha, quaha quaha quaha'
    • 10-18 calls produced in 5 seconds
    • Barks linked by an inhaled 'ee' sound
• Snort
  • Short (c. 1 second) snorts indicate agitation
  • Long (> 2 seconds) snorts express well-being, as when feeding
• Vocal expressions with unknown significance
  • 'ee-ha' - a two-syllable sound
  • High-pitched squeal
  • Long squeaking
  • Blowing with loose lips; typically while grazing (Estes 2012)

Facial expression (from Klingel 2013 unless otherwise noted)

• 6 basic expressions
  • Reflect friendly or curious intentions, sexual receptivity, submission, or aggression
• Friendly intent
  • Naso-nasal rubbing, also known as an "empty grooming gesture" (see description above)
  • Flehmen, a curling of the lips, is a reaction to an interesting odor
    • Head held up
    • Upper lip pulled back to expose the incisors
    • Lower lip pulled down
    • Jaw remains closed
• Sexual receptivity
  • Estrus face (description here)
• Submissive displays
  • Similar to estrus face, though jaw movements are faster and the chewing teeth often meet one another
• Aggressive expressions
  • Ears held slightly back with lower intensity aggression
  • Head lowered with neck outstretched, ears nearly strait back, teeth bared
    • Body sways from side to side in the highest intensity situations
    • Demonstrates readiness to fight, especially to bite
    • Recipient of display often flees or readies for an attack
• Expressions with unknown significance
  • Lip clapping
  • Yawning

Olfaction/Scent Marking (from Estes 2012 unless otherwise noted)

• Dunging ritual
  • Performed by stallions
  • Dung of herd mares or rival males covered with own feces
• Urine of mares sniffed by stallions
  • Inspection typically includes flehmen (lip curl)

Walk, trot, gallop, and run

• Gaits the same as those naturally occurring in horses (Estes 2012)
• Built for speed and long distance travel (Estes 2012)
  • Maximum speed: 60-70 km/hr (37-43 mi/hr) (Grubb 1981; Kingdon 1979)

Swim to cross rivers and streams

• Leg movements
• Resemble that of a trotting horse (Waring 1983)
• Head elevated to place eyes and nostrils above the water (Waring 1983)

Other Behaviors

• Dust bath (Estes 2012; Waring 1983)
  • Roll onto the back in the dust
    • Capable of rolling completely over from one side to the other
  • All four legs often held skyward as the zebra wiggles from side to side
• Rub head and body against objects (Estes 2012)
  • Trees, rocks, and termite mounds used to scratch
• Groom (from Waring 1983 unless otherwise noted)
  • Nibble and scrape hair with incisors
• Tail switch (from Waring 1983 unless otherwise noted)
  • Flick tail to strike the body sides
  • Often to discourages biting insects

Predators

• Major predators (Grubb 1981; Klingel 2013)
  • Lion (Panthera leo) and spotted hyena (Crocuta crocuta)
    • May take adult zebra (Grubb 1981)
    • Sizeable hyena pack typically required to capture a zebra
  • Cheetah (Acinonyx jubatus), African wild dog (Lycaon pictus), and leopard (Panthera pardus)
    • Primarily target foals (Grubb 1981)
  • Nile crocodile (Crocodylus niloticus) opportunistically take zebra
• Antipredator behaviors (from Estes 2012 unless otherwise noted)
  • Harem members often form tight associations and stand their ground
    • Young protected by mother and family members
      • Mother and young hide behind fellow harem members
    • Active defense by the stallion
    • Mother mounts attack if her foal is captured (Klingel 2013)
  • Attacks made with teeth and hooves (Klingel 2013)
  • Strategies highly successful against many predators (Estes 2012; Klingel 2013)

Herbivore interactions

• Zebras may open up grassland habitat for use by others
  • Removal of lower-quality, older-growth grass stems, sheaths, and seed heads by zebra clears the way for more selective ruminants (Hack et al. 2002; Maddock 1979; Owaga 1975)
    • Wildebeest and Thomson's often move into migratory areas after zebra (Maddock 1979)

Symbiotic interactions

• Proposed mutualistic interactions
  • Oxpeckers (Buphagus africanus and B. erythrorynchus) and Pied Crow (Corcus albus)
    • Stand on and glean ticks from ungulates (Bishop and Bishop 2013; Breitwisch 1992)
    • Relationship not fully mutualistic; oxpeckers often wound-feed from hosts (Grobler and Charsley 1978; Nunn et al. 2011; Plantan et al. 2013; Weeks 2000)
    • Controlled study suggests oxpeckers minimally reduce adult tick load and prolong healing time of wounds (Weeks 2000)
  • Ostrich (Struthio camelus) feed in association with many ungulates (Dean and MacDonald 1981; Perry 1983)
    • Speculation that partners benefit from one another's abilities to detect predators
    • Few studies conducted, none with data to support mutualism
• Commensalistic interactions (Dean and MacDonald 1981)
  • Some birds eat insects flushed out by walking ungulates; most do not exclusively feed in this manner
    • Cattle egret (Ardeola ibis), Carmine Bee-eater (Merops nubicus), Wattled Starling (Creatophora cinerae), Fork-tailed Drongo (Dicrurus adsimilis), Piapiac (Ptilostomus afer)
    • Often seen on the back of ungulates or standing beside them
• Ostrich (Struthio camelus) feed on beetles attracted to zebra dung (Dean and MacDonald 1981)
• Parasitic interactions (from Klingel 2013 unless otherwise noted)
  • Healthy individuals often carry a high parasitic burden
    • Ill effects likely manifest only in individuals already weakened or stressed
  • Numerous nematodes, cestodes, and trematodes
  • Ectoparasites include flees, lice, and ticks
    • Fleas inflict bites and lay eggs in the hair
      • Ingested larvae attach within the intestinal tract to feed and complete their life cycle

Aggressive displays are similar to those observed in the domestic horses. Individuals strike out with hooves, neck wrestle, bite and chase their opponents.

Image credit: © P Steward from Flickr. Some rights reserved.

Plains zebra are highly social animals that live in small family units that maintain contact with one another even when mixing into larger herds of ungulates containing several thousand individuals. Close family bonds are maintained through social grooming, close standing, and greeting ceremonies. This picture, taken in Namibia, shows group-mates as they rest their heads on one another's back.

Image credit: © J Oldenettel from Flickr. Some rights reserved

Bishop and Bishop (2013)
Breitwisch (1992)
Dean and MacDonald (1981)
Estes (2012)
Grobler and Charsley (1978)
Grubb (1981)
Hack et al. (2002)
Harris et al. (2009)
Kingdon (1979)
Klingel (1965)
Klingel (1967)
Klingel (1969)
Klingel (2013)
Maddock (1979)
Naidoo et al. (2014)
Nunn et al. (2011)
Owaga (1975)
Perry (1983)
Plantan et al. (2013)
Schilder (1988)
Smuts (1975)
Smuts (1976c)
Waring (1983)
Weeks (2000)