Myth, Marula, and Elephant: An Assessment of Voluntary Ethanol
Intoxication of the African Elephant (Loxodonta africana)
Following Feeding on the Fruit of the Marula Tree
School of Biological Sciences, University of Bristol,
Woodland Road, Bristol BS8 1UG, United Kingdom
Accepted 6/1/2005; Electronically Published 2/6/2006
Africa can stir wild and fanciful notions in the casual visitor;
one of these is the tale of inebriated wild elephants. The sug-
gestion that the African elephant (Loxodonta africana)becomes
intoxicated from eating the fruit of the marula tree (Sclerocarya
birrea) is an attractive, established, and persistent tale.Thisidea
now permeates the African tourist industry, historical travel-
ogues, the popular press, and even scholastic works. Accounts
of ethanol inebriation in animals under natural conditions ap-
pear mired in folklore. Elephants are attracted to alcohol, but
there is no clear evidence of inebriation in the field. Extrap-
olating from human physiology, a 3,000-kg elephant would
require the ingestion of between 10 and 27 L of 7% ethanol
in a short period to overtly affect behavior, which is unlikely
in the wild. Interpolating from ecological circumstances and
assuming rather unrealistically that marula fruit contain 3%
ethanol, an elephant feeding normally might attain an ethanol
dose of 0.3 g kg?1, about half that required. Physiologicalissues
to resolve include alcohol dehydrogenase activity and ethanol
clearance rates in elephants, as well as values for marula fruit
alcohol content. These models were highly biased in favor of
inebriation but even so failed to show that elephants can or-
dinarily become drunk. Such tales, it seems, may result from
“humanizing” elephant behavior.
*This paper was prepared as an overview of a symposium session presented at
“Animals and Environments,” the Third International Conference for Com-
parative Physiology and Biochemistry, Ithala Game Reserve, KwaZulu-Natal,
South Africa, 2004(http://www.natural-events.com/ithala/default-follow_2.asp).
†Corresponding author; e-mail: email@example.com.
Physiological and Biochemical Zoology 79(2):363–369. 2006. ? 2006 by The
University of Chicago. All rights reserved. 1522-2152/2006/7902-5055$15.00
Africa is intoxicatinginitsvistasandwildnessandinitsvarieties
of bottled liqueurs. First-time visitors frequently imagine rav-
enous beasts behind each bush, a fancy brought on by the
undiluted and visceral attraction of wild Africa and, perhaps,
by an indulgence with the liqueurs. Commercial brewing and
distilling are important industries in South Africa, and among
the more popular liqueurs is Amarula, made from the fruit of
the marula tree, whose bottle is adorned with marula fruit and
a rampant bull elephant. (The producers of Amarula [Distell
Group Ltd., Stellenbosch, South Africa] make no claim as to
marula intoxication of elephants but correctlydescribethelocal
folklore that elephants are insatiably attracted to the marula
tree, known locally as the “elephant tree”; http://www
ference for Comparative Physiology and Biochemistry, in the
Ithala Game Reserve, South Africa, we were prompted toreflect
on these issues by the persistently related anecdote of drunken
elephants. Even a superficial perusal of promotional material
from the African safari and tourist industry reveals a widely
held belief that evidence exists for ethanol intoxication in the
African elephant and most especially that this elephant (Lox-
odonta africana) becomes intoxicated from eating the ripe and
fermenting fruit of the marula tree (Sclerocarya birrea). This
belief is persistent. For example, the French naturalist Dele-
gorgue, describing his experiences in thevicinityoftheiMfolozi
River in present-day KwaZulu-Natal around 1839, reports how
his Zulu guides told him of the strangely aggressive behavior
shown by elephant bulls after feeding on marula fruits. He
writes, “The elephant has in common with man a predilection
for a gentle warming of the brain induced by fruit which has
been fermented by the action of the sun” (Alexander and Webb
1990; p. 275). Naturally, at a meeting with physiologists, there
were some attempts to calculate whether this was a likely phe-
nomenon, most befuddled by attempts at empirical proof with
the Amarula. In this paper, we explore and analyze the likeli-
hood of natural intoxication in the African savannah elephant.
A recent symposium at the Society for Integrative and Com-
Elephant and Marula
Okagbue R.N. and M. Siwela. 2002. Yeasts and related micro-
organisms isolated from ripe marula fruits (Sclerocarya caf-
fra) in Zimbabwe. S Afr J Sci 98:551–552.
Owen-Smith R.N. 1988. Megaherbivores: The Influenceof Very
Large Body Size on Ecology. Cambridge University Press,
Poole J. 1997. The Elephants. 2nd ed. Colin Baxter Photog-
Prinzinger R. and G.A. Hakimi. 1996. Alcohol resorption and
J Ornithol 137:319–327.
Rees P.A. 1982. Gross assimilation, efficiency and food passage
time in the African elephant. Afr J Ecol 20:193–198.
Sa ´nchez F.,C.Korine, B.Pinshow,andR.Dudley.2004.Possible
roles of ethanol in the relationship between plants and fru-
givores: first experiments with Egyptian fruit bats. Integr
Comp Biol 44:290–294.
Shackleton C. 2002. Growth and fruit production of Sclerocarya
birrea in the South African lowveld. Agrofor Syst 55:175–180.
Shoshani J. 1992. Elephants: Majestic Creatures of the Wild.
Rodale, Emmaus, PA.
Siegel R.K. 1989. Intoxication: Life in Pursuit of Artificial Par-
adise. Dutton, New York.
Siegel R.K. and M. Brodie. 1984. Alcohol self-administration
by elephants. Bull Psychon Soc 22:49–52.
Spinage C. 1994. Elephants. Poyser, London.
Steenkamp V. 2003. Traditional herbal remedies used by South
African women for gynaecological complaints. J Ethnophar-
Sukumar R. 2003. The Living Elephants: Evolutionary Ecology,
Behaviour and Conservation. Oxford University Press,
Tang A.M.C., K.D. Hyde, and R.T. Corlett. 2003. Diversity of
fungi on wild fruits in Hong Kong. Fungal Divers 14:165–
West L.J., C.M. Pierce, and W.D. Thomas. 1962. Lysergic acid
diethylamide: its effect on a male Asiatic elephant. Science
White L.T.J., C.E.G. Turin, and M. Fernandez. 1993. Group
composition and diet of forest elephants, Loxodonta africana
cyclotis Matschie 1900, in the Lope ´ Reserve, Gabon. Afr J
Wolf F.W. and U. Heberlein. 2003. Invertebrate models of drug
abuse. J Neurobiol 54:161–178.
Wyatt J.R. and S.K. Eltringham. 1974. The daily activity of the
elephant in the Rwenzori National Park Uganda. E Afr Wildl
Yoon C.K. 2004. “Of drunken elephants, tipsy fish and scotch
with a twist.” New York Times, March 23, late edition–final,