The Sounds of Africa – Lions

The Lion Roar is the Voice and Sound of Africa

Off in the distance in the savannahs of Africa, you listen, and a roar comes out of the deep, creating an uneasy trepidation.  The lion’s roar is indeed the voice and sound of Africa.  It sends an awesome message to the world that he is lord and master to all he surveys.

For a human observer, the roaring of a lion – even more so that of a whole pride – is one of the most impressive vocalizations in the animal kingdom.  The author videoed multiple lion experiences in the Mala Mala and Sabi Sands areas of Mpumalanga, South Africa.  The following was a rather “interesting” experience with lions at close range one evening when stuck in the Sabi River bed in Mala Mala South Africa, with a pride of 17 lions surrounding the vehicle.  The roars are about 2/3^rd through this first video. Fortunately, the lion pride had already eaten for the evening.

Unlike most large cats, lions are very social animals that live in prides.  A pride can be composed of up to thirty lions, although most are smaller.  The pride contains one or more males, who are often brothers, and related adult females and their cubs.  Each pride has its own hierarchical system, with one dominant male as the master of the pride and territory.  The weaker males rank above all females.  Both males and females hunt for the pride and help defend against predators.

However, living in such close contact requires that lions have very effective communication to live together successfully.  This level of communication is greater for lions than for other large cats (leopards, cheetahs), which are more antisocial than lions.  As a result, lions produce a greater variety of sounds.  Still, the language of most cats is less developed and complicated than that of other animals.

Lions are social animals, so after all that sleeping and relaxation (up to 20 hours a day), hunting and eating, the other important aspect of the lion’s life is communicating.

Lions communicate with one another through a variety of methods, including body language, appearance, behavior, touch, scent, and vocalization.

1. Social order.  Dominant males are most apt to let a pride member know when he or she is out of order.  This is often done with a quick swat of the paw or a gentle bite to the neck.  They also communicate by example, as in a staged hunt where cubs are shown how the hunt is done.  And, lions also show affection in their own special way when relaxing by licking gently or gnawing at each other.
2. Appearance.  The impressive mane of the male gives the appearance of a large head, and therefore the impression of a significant threat.  The male’s size and attitude of eerie calm are often sufficient to let any upstarts know to back off.  But, this is not their only signaling tool.  The large teeth of the lion are certainly functional, but they also serve to let others know just how powerful their bite can be.  Baring teeth essentially says “don’t mess with me.”
3. Behavior.  Lions also communicate with lions from outside the pride that attempt to compete with them for resources or territory.  They use urine to delineate the borders of their territory; the scent warns others that crossing the line means certain conflict.  Lions also form coalitions, often made up of male relatives, to patrol the claimed areas.  They stand at the borders and stare down those who have ventured too close for comfort.  If staring isn’t enough to scare off the intruders, a lion will let out a powerful roar.  This signals that the lion means business – an indication of his strength and intention to fight.
4. Touch.  Lions from the same pride greet each other by rubbing their cheeks, necks, and bodies together.  Grooming each other helps the pride bond with each other.  Their hard bumpy tongues are used to comb each other’s fur and remove ticks.  When lions are relaxing they show affection by licking gently or gnawing at each other.
5. Scent.  Lions have many scent glands on their cheeks, lips, forehead, chin, tail, and even between their toes.  Scent communicates with other lions where they have been and how long ago.  Scent is also used to attract a mate.  They can smell other cats using an extra smell organ called the Jacobson’s organ.  To use this organ, the lion shuts off its nose, curls its upper lip, and sucks air through its mouth.  Males also leave a calling card in the area consisting of a scent mixed with urine on bushes and turfs of grass (Animal Behavior p72). This signals to any nomadic lion that he is in another lion’s territory.
6. Posturing.  Lions use many parts of their bodies to communicate their feelings, with the most important part being their face.  When they feel threatened and scared, they show their weapons (claws and teeth), stand on their tiptoes, raise their tails, and hunch their backs in an attempt to look at large as possible.  The dominant male will walk with his head held high, ears standing upright, mouth closed, and tail swishing side-to-side.  The more submissive members of the pride walk with their heads held low, eyes flattened against their heads, mouths open, and teeth showing.  It is said that lions are able to detect event the slightest change in posture of another lion.
7. Vocalizations.  Lions communicate with different noises, such as ferocious roars to call another lion or a soft rumble to indicate their satisfaction.

The Roar of the Lion

In the pride, both sexes roar to broadcast ownership and defense of a territory, or under some circumstances, to attract mates. Male lions roar to advertise their territory to other lions, whereas female lions roar to maintain contact with other lionesses, protection, or location.

Lionesses can also estimate the number of individuals roaring, and are less likely to approach foreign roars when they are outnumbered. A second video shows the result of a male lion in a tree, having taken a leopard’s impala kill, dropping to the ground, and stopping a very short distance from us to eat the remains of the carcass.  Listen to the sounds of the feast.

An additional function might also be coordination of hunting where they can communicate differently other than with a roar.  On the hunt a lioness lashes her tail, but just before the final attack,  holds it straight out behind. To the lioness the roar means protection or just “Here I am.”  If enemies approach a lioness, she will give off warning signals such as flattering her eyelids and lashing her tail.  Females and their cubs use the largest variety of calls to communicate because they are the members of the pride that spend the most time together.

Nomadic males wander widely, passing through pride ranges single or in coalitions until they are successful in taking over a pride of their own.  Roaring provides a means by which nomadic males might co-ordinate their movements with coalition partners or to recruit new ones.

However, if nomads used this loud long-distance signal to communicate with social companions they would also advertise their position to resident males in the area.  The costs to nomadic males on engaging the attention of resident males may be considerable.  Resident male lions consistently give aggressive approaches to roaring from strange males broadcast in their territories.  Observations indicate that real intercoalition encounters may be fatal.  Interestingly, nomad lions without a pride are unlikely to roar in another lion’s territory unless a challenge to the pride control is being made.

Roar Described

Roars are loud, low-pitched vocalizations that are delivered in bouts, which typically last 30 to 60 seconds and consist of several soft introductory moans, a series of full-throated roars and a terminating sequence of grunts.   Cooperation in male lions: kinship, reciprocity or mutualism? Lions are active primarily at night, and most roaring occurs during nocturnal hours .

 

To be even more specific, lion roaring has two distinct physiological and acoustic components:

1. A low fundamental frequency, made possible by long or heavy vocal folds, which lead to the low pitch of the roar;
2. Lowered formant frequencies, made possible by an elongated vocal tract, which provide the impressive baritone timbre of roars.

The lion is the loudest of all the big cats with their roar reaching up to 114 dB SPL at 1 meter.*  Because they are one of the most threatening predators in the wild, they are not concerned about being too noisy.  Lions are active primarily at night, and most roaring occurs during nocturnal hours.

Spectrographic analysis of lion roar reveals both low-frequency components with a fundamental frequency around 180-194 Hz and a higher-frequency component around 4,000 Hz.  A gender difference also exists with the fundamental frequency for males at about 195 Hz and for females about 207 Hz.  The roar can be heard at least 5 miles (8 km).

The lion roar is a series of calls with a fairly regular structure consisting of a single call followed by a series.  The series, however, changes in call type, intensity, temporal sequencing, duration, and intervals between them.  A typical lion roaring can last for more than a minute, usually starting with a few low-intensity moan-like calls, then progressively increasing in intensity and duration, and upon approaching the intensity climax the calls become shorter again and harsher.  Following the climax is a series of short harsh calls, in the beginning uttered at fairly monotonic intensity and brief intervals between the calls, then towards the end of the series gradually decreasing in intensity and with increasing interval duration.

A third video shows a pride of lions on an impala kill at night.  Listen to the sounds of the pride while dining.

*I have added the SPL identifier to the sound level because the reference did not list it as such.  However, this seems like a reasonable assumption because sound level meters seem to have been the measurement tool used.

 

Next up – the Leopard

 

References:

1. Schaller, G. B. 1972. The Serengeti Lion. Chicago: University of Chicago Press
2. Lynn, R.  Life in the pride.  Disney Adventures, July 30, 1994, p29-31
3. McComb, K., Packer, C. & Pusey, A. 1994. Roaring and numerical assessment in contests between groups of female lions, Panthera leo. Animal Behaviour, 47, 379–387
4. Hanby, J. P. & Bygott, J. D. 1987. Why do subadult lions leave their natal pride? Animal Behaviour, 35, 161–169
5. Pusey, A. E. & Packer, C. 1987. The evolution of sex-biased dispersal in lions. Behaviour, 101, 275–310
6. Grinnell, J., Packer, C. & Pusey, A. E. 1995. Cooperation in male lions: kinship, reciprocity or mutualism? Animal Behaviour, 49, 95–105
7. Weissengruber, G. E., Forstenpointer, G., Peters, G., Kubber-Heiss, A. and Fitch, W. T. (2002). Hyoid apparatus and pharynx in the lion (Pantera leo), jaguar (Panthera onca), tiger (Pantera tigris), cheetah (Acinonyx jubatus), and domestic cat (Felis silvestris f. catus). J. Anat. 201, 195-209
8. Eklund, R., Peters, G., and Mabiza, E.  An acoustic analysis of lion roars. II: vocal tract characteristics.  2011, TMH-QPSR Vol. 51, p 5-8
9. Sunquist, M. and Sunquist, F. Wild Cats of the World. 2002.  University of Chicago Press, 462 pages

 

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Wayne Staab, PhD, is an internationally recognized authority in hearing aids. As President of Dr. Wayne J. Staab and Associates, he is engaged in consulting, research, development, manufacturing, education, and marketing projects related to hearing. His professional career has included University teaching, hearing clinic work, hearing aid company management and sales, and extensive work with engineering in developing and bringing new technology and products to the discipline of hearing. This varied background allows him to couple manufacturing and business with the science of acoustics to bring innovative developments and insights to our discipline. Dr. Staab has authored numerous books, chapters, and articles related to hearing aids and their fitting, and is an internationally-requested presenter. He is a past President and past Executive Director of the American Auditory Society and a retired Fellow of the International Collegium of Rehabilitative Audiology. 

 

_{**this piece has been updated for clarity. It originally published on October 16, 2012}