Spotted Hyena (Crocuta crocuta) Fact Sheet: Behavior & Ecology

Daily Activity Patterns

• Nocturnal most often
• Active at night, though some daylight activity is not uncommon (Kruuk 1972)
  • Hunt at night, possibly to avoid daytime heat (Cooper 1990; Mills 1984b)
  • 2 peaks in activity occur, near mid-night and prior to sunrise; similar pattern commonly observed in other nocturnal animals (Kruuk 1972)
  • Most movement between sunset and shortly after sunrise (Stratford and Stratford 2011)
    • Found alone or in pairs most often (Hofer and East 1993b; Kruuk 1972)
    • Move long distances each night, especially when water and food resources are limited (Hofer and East 1993b; Stratford and Stratford 2011)
    • Travel c. 27 km/night, as measured in one population in the southern Kalahari (Mills 1984b)
    • Maximum distances of 80 km (see summary table in Hofer and East 1993b)
• Rest for long periods during the day and night (Kruuk 1972; Stratford and Stratford 2011)
  • c. 50-80% of time spent resting/not moving, though more active during the wet season in another study (Kruuk 1972; Stratford and Stratford 2011)
  • Females with cubs often rest at the den site
  • Individuals seek cool, shady spots to rest during the day; lie in the open in cooler weather
    • Rest inside holes (not in dens), at lake and stream edges in the mud, or inside dense shrub
• Nursing cubs are typically active at dawn and dusk (Hofer and East 1993c)

Annual Activity Patterns (summarized from Hofer and East 1993b unless otherwise noted)

• Commute to take advantage of migratory herds of prey
• Serengeti clans invest large amounts of time annually to travel between clan territory and the nearest migratory herds, when prey density within territorial boundaries is low
  • May cover distances >70 km
  • c. 24% of all commutes are > 40 km
  • Denning females (those with dependent cubs) may spend nearly 50% of the year commuting between clan territory and the nearest herds of migratory herbivores (Hofer and East 1993c)

Territory features

• Territory function and use
• Serve as breeding and feeding areas (Hayward et al. 2009; Hofer and East 1993a)
• Foraging trips outside of territorial boundaries are not uncommon (Honer et al. 2005)

Territory size

• Broad range of sizes reported
• 9-1000⁺ km² (East and Hofer 2013)
• Size likely related to prey availability (Hayward et al. 2009; Hofer and East 1993a)
• Smaller territories typical for clans living in grassland habitats which support greater prey biomass
  • c. 32 km2; Aberdare National Park, Kenya - small (70 km²) mosaic forest habitat (Sillero-Zubiri and Gotteli 1992a)
  • c. 60 km²; Masai Mara National Reserve, Kenya - predominantly rolling grasslands (Boydston et al. 2003)
  • c. 91 km²; Addo Elephant National park, southern Africa - predominantly thicket with some grassland (Hayward et al. 2009)
  • c. 570 km²; Namib-Naukluft Park, Namibia - region of the Namibian desert along the Kuiseb River (Tilson and Henschel 1986)

Highly social

• Female dominated society
• One of the most social of all large carnivores
• Form stable social units, though members may spend much time away from clan mates (Frank 1986b; Hofer and East 1993a; Kruuk 1972)
  • Described as a "fission-fusion" society (East and Hofer 2013)
    • Subsets of the clan forage for prey (Cooper 1990; Mills 1984b)

Live in clans

• Clan composition
• Composed of one to several matrilineal lines and immigrant males (East and Hofer 2013)
• Sex ratio typically evenly distributed between males and females (Hofer and East 1993a)
• All adult females are reproductively active (Hofer and East 1993a; East and Hofer 2001)
• Male members remain with their chosen clan for many years (Frank 1986b; Hofer and East 1993a; Kruuk 1972)
• Females often remain in their natal clan for life (Frank 1986b; Hofer and East 1993a; Kruuk 1972)
• Cooperate to defend territory, food resources, and the communal den (East and Hofer 2013)

Clan size (Cooper et al. 1999; Frank 1986b; Kruuk 1972; Mills 1984b)

• Range: 8-80 individuals
• Varies across distribution
• Smaller clans in regions with limited food resources (Mills 1984b)
  • Eg. the southern Kalahari where clans of c. 8 individuals are reported (Mills 1984b)
• Larger clans in areas with high prey density or seasonal spikes in prey density due to an influx of migratory herbivores (Hofer and East 1993a; Hofer and East 1993b)
  • Eg. the Serengeti where clans of c. 55-75 individuals are reported (Hofer and East 1993a; Hofer and East 1993b)

Social hierarchy

• Benefits of social rank
• Rank determines access to food and other critical resources (Frank 1986b; Kruuk 1972)
  • Age and body size do not dictate social rank for males or females (East and Hofer 2001; Frank 1986b; Holekamp et al. 1996)
• Higher ranking females experience greater reproductive success (Holekamp et al. 1996)
• Dominance hierarchies
• Females dominate all males (Frank 1986b; Kruuk 1972)
• Linear dominance hierarchy among clan females (East and Hofer 2013; Frank 1986b)
• Males form a separate linear hierarchy among themselves (East and Hofer 2013; Frank 1986b)
• Features of the female social hierarchy
  • Mother's social status dictates social rank of her offspring (Frank 1986b; Kruuk 1972)
    • Offspring receive a rank immediately below that of the mother upon reaching maturity (East and Hofer 2013)
    • Likely the result of learned behavioral mechanisms; offspring mimic the social interactions of the mother, with maternal support (East and Hofer 2013)
      • Adopted cubs receive ranks of adoptive mothers, not birth mothers (East and Hofer 2013)
  • Reproductive success influenced by social status
    • Dominant females have greater reproductive success than lower ranking counterparts; due to the combined effects of lower age at first reproduction, increased ability to overlap lactation and pregnancy, shorter intervals between consecutive births, and higher rates of offspring survival (Holekamp et al. 1996)
    • Low ranking females more impacted by prey scarcity (Holekamp et al. 1996)
• Features of the male social hierarchy
  • Social rank gained by strict adherence to "queuing" conventions (East and Hofer 2001; East et al. 2003)
    • Clan tenure determines rank within the queue; social dominance gradually increases through time (East and Hofer 2001)
    • Time to gain "alpha" status is positively related to clan size (East and Hofer 2001)
    • Long tenured males form coalitions securing their high rank and mating privileges with older, highly ranked females (East and Hofer 2001)
    • Coalitions cooperatively "guard" older females from sexual advances by subordinate males (East and Hofer 2001)
    • Non-dispersing males are the exception; they receive top rank quickly upon reaching sexual maturity (East and Hofer 2001)
  • Physical contests between clan males are rare and they do not appear to play a role in gaining or losing social status (East and Hofer 2001; East et al. 2003)
  • Dominance is expressed vocally by producing a loud call or whoop; the frequency and duration of whooping bouts signal rank (East and Hofer 1991b)
    • High-ranked males whoop more frequently and for longer periods of time than lower-ranked males
  • High ranking males are more likely to gain matings with high-ranking females

• Greeting "ceremonies"
  • Unlike those of the brown hyena (Kruuk 1972; Mills 1984b)
  • Solidify social bonds between individuals that often range broadly but come together at communal dens in care of young (Kruuk 1972)
  • Behavior performed by all clan members; irrespective of gender, age, or social status (Mills 1984b)
  • Display description
  • One individual approaches the other, often with great caution
  • Briefly sniff at one another's side, mouth, head, and neck before standing parallel with heads in opposite directions (Kruuk 1972)
  • Lift leg nearest one another as each mutually sniffs and licks at the other's erected genital organs (Mills 1984b)
    • No sexual significance despite the incorporation of genital sniffing (Mills 1984b)

Advertise to claim territory

• Call, scent-mark, and regularly patrol (East and Hofer 2013)
• Border patrols regularly carried out by adult clan members (Boydston et al. 2001)
  • All patrolling members scent-mark (see description of olfaction/scent marking below)
  • Lead by females most often (Boydston et al. 2001)
  • 2 or more individuals often walk in parallel (Kruuk 1972)

Intruders

• Most commonly non-resident males (Boydston et al. 2001)
  
• Reponses to intruders (Boydston et al. 2001; Hofer and East 1993b)
• Commonly left alone or given a cursory investigation (eg. sniffing); non-residents often demonstrate signs of submission or retreat before residents move toward them (Hofer and East 1993b)
• Chased for a distance before being left or aggressively engaged
• Neutral outcomes often in one-on-one encounters and in cases where non-residents are clearly in transit through the residents' territory (Hofer and East 1993b)
• "Parallel walk", a visual threat to an intruder
  • Joint display, performed by 2 or more individuals (Frank et al. 1989; Kruuk 1972)

Interclan fighting

• Physical contact is rare (Kruuk 1972)
• Most "fights" involve vocalization, body posturing, and chasing
• Adult females are more likely to initiate/lead fights than are adult males (Boydston et al. 2001)
• Physical aggression is often deadly (Kruuk 1972)
• Fight frequency
  • Highly variable across populations (Boydston et al. 2001)
  • Encounters at kills made close to territorial boundaries often escalate into fights (Boydston et al. 2001; Hofer and East 1993a; Hofer and East 1993b)

• Gender specific aggressive responses (Boydston et al. 2001)
  • Resident females tend to attack female intruders more than male intruders
  • Resident males tend to attack male intruders more than female intruders
  • Most attacks on intruding males are initiated by resident males

Aggressive displays (from East and Hofer 2001 unless otherwise noted)

• Intraclan (within clan) aggression generally rare
  • Display and posturing reduce occurance of physical encounters
• Threat displays
  • Hair over the back and neck held erect; as with other hyenas (Mills 1984b)
• Gender and aggression
  • Females most commonly initiate aggressive behavior (East and Hofer 1991b)
  • Aggression between males does not dictate social status
  • Females are rarely the target of male aggressive acts
  • c. 6% of observed interactions in one study
  • Most often by mid-ranked males lunging and biting females, sometimes causing injury to the leg or rump
• Forms of physical aggression
  
  • Lunge, push, chase, bite, wrestle, and pin opponents to the ground (East and Hofer 1991b; East and Hofer 2001; Kruuk 1972)
  • Teeth, not claws, used as weapons during fights (East and Hofer 1991b; East and Hofer 2001; Kruuk 1972)

Submissive displays (from East and Hofer 2001)

• Subordinates display to avoid physical fights
• Retreat or are displaced
• Cower; tail held between the legs, ears back, head bobbing or held upside down

Participants (from Kruuk 1972)

• Common among cubs and young
• Often chase and bite at one another
• Not unfrequently among adults
• Often play along with young

Forms of Play (from Kruuk 1972)

• Adult play appears similar to some human forms
• "Keep Away" - circular chasing, often near large carcasses while eating
  • A hyena with a piece of meat or bone often chased by another, no apparent attempt is made to evade his pursuer
  • Participants often trade roles
• Splashing and dunking one another while swimming

Vocal features (from Kruuk 1972 unless otherwise noted)

• Largest vocal repertoire of all hyena species
• Frequently vocalize
• Vocalize individually most often, unlike other social carnivores (eg. wolves, Canis lupus), which commonly vocalize as a group (East and Hofer 1991b)
• Individual voice recognition possible
• Acoustic signatures present within many calls (East and Hofer 1991a; East and Hofer 1991b)

Long distance call, the whoop (from East and Hofer 1991b unless otherwise noted)

• Functions
• Display identity, designate location, or request support
• Near den sites, the significance of adult calls appears gender dependent
  • Males whoop more frequently than females
    • Dominant males tend to direct calls toward females
    • Subordinate males tend to direct calls toward other males
  • Female whoop often as vocal defense of the denning area or food resources
  • Alpha-males and alpha-females vocalize more than other clan members of the same sex
• Auditory characteristics
• Audible by humans >5 km (Kruuk 1972; Mills 1984b; East and Hofer 1991a)
• Often in the form of whooping bouts, composed of a number of discrete whoop calls
  • Given at a den site or when approaching a carcass
• Performed while walking, loping, standing, or lying (Kruuk 1972; Mills 1984b; East and Hofer 1991a)
• See Kruuk 1972 for descriptions of body posture, auditory characteristics, and contextual occurrence
• Call variation
• 3 variations (Type A, S, and T), characterized by the degree of symmetry throughout the call and variation in sound frequency/pitch (East and Hofer 1991a)
  • Type A and S calls begin with extended low frequency production before rising in pitch and returning to the initial frequency
  • Type T calls lack variation in pitch
• Cubs often whoop (East and Hofer 1991a; East and Hofer 1991b)
  • Commonly after being attacked or aggressively threatened (East and Hofer 1991a; East and Hofer 1991b)
  • Occasionally to call to or respond to their mothers (East and Hofer 1991a; East and Hofer 1991b)
• Inter-clan whooping occurs, though at low frequency
  • Vocal exchanges may last for nearly an hour, with whooping bouts c. every 2 minutes within a single clan

Close range calls (from Kruuk 1972; Mills 1989)

• Commonly exchanged in communal antagonistic situations
• E.g. in defense of food and territory, or against lions
• Groan, giggle, yell, growl, grunt, low, and squeal

Cub vocalization (from Kruuk 1972)

• Whine
• Associated with feeding situations
• To initiate suckling or at a carcass when feeding attempts are thwarted
• Soft squeal
• A form of greeting

Function of scent marking

• Primary form of intraspecific communication between clans (Kruuk 1972; Mills 1984b)

Forms of scent marking

• Defecation
  • Maintain communal latrine sites (Bearder and Randall 1978; Kruuk 1972)
    • Often scattered throughout the home range along heavily travelled routes or other conspicuous locations (Bearder and Randall 1978)
    • Often near territory boundaries (Hofer and East 1993a; Kruuk 1972)
  • Evidence for prolonged use of the same site; several years in some cases (Bearder and Randall 1978)
    
    • Whitish colored feces often covering 10 m in diameter (Kruuk 1972)
    • Calcium-rich feces decompose slowly; remain visible for up to 14 months (Bearder and Randall 1978)
• Pasting
  
  • A unique hyenid habit (Mills 1989)
  • Strong-smelling, whitish, fatty secretions from the anal gland deposited onto grass stalks or shrubs (Mills 1989)
    • A semi-crouched position is taken as the individual slowly walks over the plant, bending the stalk and passing it under the body, between the fore- and hind legs; typically repeated several times (Kruuk 1972)
    • Gland is partially protruded to release secretions (Kruuk 1972)
  • Commonly performed while walking alone, around kills, when lions are present near dens, by groups patrolling near territorial borders (Kruuk 1972)
  • May be aggressively motivated (Kruuk 1972; Mills 1984b)
• Scratching
  • Interdigital glands on the feet commonly used to scent mark following pasting or defecation (Kruuk 1972)
    • No attempt to cover feces, as with domestic cats
  • Often observed when males are sexually aroused (East and Hofer 1991b)
  • Females rarely mark in this way (East and Hofer 1991b)

Run

• A loping gate (East and Hofer 2013)
• Speed
• Travel c. 10 km/hr (c. 6.2 mi/hr) (Mills 1984b)
• Exceed 50 km/hr (31 mi/hr) in pursuit of prey (Kruuk 1972; Mills 1984b)
• Tail held strait behind body (Kruuk 1972; Mills 1984b)

Swim (from Kruuk 1972)

• Strong swimmers
• Venture into water for a meal or to reach suitably shaded resting grounds
• Observed swimming at least 200 m distance
• Dive to tear flesh from an animal drowned in a lake
• Submerging for up to 12 seconds
• Smooth, dog-like style

Non-arboreal

• Do not climb trees (Kruuk 1972)

Carnivore competition (from Kruuk 1972 unless otherwise noted)

• Numerous potential competitors
  
  • Black-backed jackal, cheetah, leopard, lion, and wild dog inhabit the territories of many spotted hyena (Bearder 1977)
  • All share a high degree of overlap in prey preference
• Lions often dominate interactions with spotted hyena
  • Hyena are often seen following lions
  • Confrontation limited, during infrequent interactions lions typically ignore hyena unless harassed
  • Solitary lions can be overtaken by a large hyena group
    • Similar to mobbing behavior displayed by some small birds toward a larger predator

Kleptoparasitism and competition for carrion

• Oportunistically steal food from others
• Stolen food accounts for a minor portion of spotted hyena annual diet, > 1%, one study  (Cooper et al. 1999)
• Associations between lions and spotted hyena are common
• These species often steal kills from one another (Cooper et al. 1999; Kruuk 1972)
  • Nature of theft influenced by predator population density and prey composition and density (Kruuk 1972)
  • Lions commonly overtake and consume hyena kills near Ngorongoro Crater, Tanzania (Kruuk 1972)
    • Over 80% of observed lion feedings are hyena kills; though more uncommon at other locations (Kruuk 1972)
• Spotted hyena displace smaller predators
• African wild dog, brown hyena, cheetah, and jackal  often lose competitions for carrion (Cooper 1990; Cooper et al. 1999)
  • E.g. brown hyena grabbed by the neck and shaken by a spotted hyena to claim a meal (Mills 1984b)
• Vultures and hyenas
• Often feed simultaneously from a carcass, especially during daylight (Kruuk 1972)
• Interactions with humans
  • Spotted hyena prey on domestic livestock and humans (though uncommonly), in some regions (Kruuk 1972)
    • Some tribes place human corpses into a bush for hyenas to dispense of the remains (eg. customary for Masai living around the Serengeti National Park and in Ngorongoro) (Kruuk 1972)

Symbiotic relationships

• Fermentative bacteria (from Theis et al. 2013)
• May contribute to species-specific odors
  
  • Bacterial communities within scent glands contribute to odor profiles of anal gland secretions
  • Composition of bacterial communities varies between species and by gender and social status within a species
• Scarab beetles (from Krell et al. 2003)
  • Dung eating and necrophagous beetles found in latrines and regurgitations
  • Prefer brownish, soft feces and regurgitation to calcium rich (white) feces

Commonly regurgitate indigestible material

• Frequently at sites where resting occurs, at dens (Bearder 1977)
• 35 regurgitations accumulating over the course of one month at one well-used den site (Bearder 1997)

Grooming (from Kruuk 1972)

• Commonly observed
• Appears similar to that observed in other carnivores
• Stretch, scratch, shake, and lick the coat and genitals
• Do not "wash" the face as with cats and some other viverrids

Spotted hyena live in socially structured clans of between 8 and 80 individuals. They spend much of their time apart. Greeting ceremonies, such as this one, act to solidify social bonds between all clan members, young and old alike.

Image credit: © D. Bygott from Flickr. Some rights reserved.

Bearder (1977)
Boydston et al. (2001)
Boydston et al. (2003)
Cooper (1990)
Cooper et al. (1999)
East and Hofer (1991a)
East and Hofer (1991b)
East and Hofer (2001)
East and Hofer (2013)
East et al. (2003)
Frank (1986b)
Frank et al. (1989)
Hayward et al. (2009)
Hofer and East (1993a)
Hofer and East (1993b)
Hofer and East (1993c)
Holekamp et al. (1996)
Honer et al. (2005)
Krell et al. (2003)
Kruuk (1972)
Mills (1984b)
Mills (1989)
Sillero-Zubiri and Gotteli (1992a)
Stratford and Stratford (2011)
Theis et al.(2013)
Tilson and Henschel (1986)