THE STUDY OF PRIMATE SELF-MEDICATION
A collection of multidisciplinary research work bymembers of The C.H.I.M.P.P. Group*(the first 18 years, 1987 – 2005)

Michael A. Huffman
Primate Research Institute, Kyoto University
Kanrin 41-2, Inuyama
Aichi Prefecture, Japan 484-8506
huffman@pri.kyoto-u.ac.jp

About this document
The following compilation of scientific research papers and articles is the result of the first18 years of fruitful collaborations from people of many nations, ideologies and cultures.
These collaborations have provide not only new insights into the wonder of nature’s
complexities, but has personally been a rewarding path to the understanding of the absolutenecessity and importance of the multidisciplinary approach and of the significance of
cultural diversity in science. Science is indeed the universal medium of our collaboration,
but no single country our discipline was more important than the others.
More than anything else, I am grateful to all who have shared in this work for their enduring
friendship and graciousness. For the results compiled here would not have been possible
had it not been for their overwhelming generosity in the sharing of valuable expertise, timeand resources. Their names are too many to mention, but the majority of them appear in thefollowing papers that make up this document.
Given the diversity of disciplines which have been involved in this research, key
collaborators from the beginning, and treasured friends, Hajime Ohigashi, Koichi
Koshimizu and I have thought for some time that is was necessary to put the growing
number of papers together into one volume, not only to provide a near complete record ofthis work, but to provide access to papers published in journals that no single reader would
be familiar with or would not normally come across in the literature of their own discipline.
Admittedly there is overlap in some of the review papers, as they were written on request or
where published as proceedings of a variety of different academic societies and congresses
inviting us to share our research with them. In spite of this overlap and the expense ofweight, I have included most of them here to provide as close of a complete record of ouractivity as possible.
There is still so much to do, as we have really only begun to scratch the surface. In
collaboration with students and colleagues from around the world, we have begun tocompile a database of the feeding lists of representative primate species ranging fromprosimians in Madagascar, monkeys of the Neotropics and Asia to the great apes of Africa
and Southeast Asia. From this we have begun to assess through the literature, and our own
research, the complimentary nutritional and medicinal properties of ‘food’ and its
importance in primates in an attempt to expand our understanding of the breadth and depthof self-medication and health maintenance behaviors and to identify promising new animaland plant targets for research. It is our strong desire that this document encourage those whoread it to pursue this line of research and to expand it above and beyond what we have done.
In the first page of each paper you will find the many institutions that have been involved
and supported each of us generously with facilities and the means of a nurturing grounds forour academic development or the support of our livelihoods. In the acknowledgements
there within you will also find the many sources of funding and facilities in the countries
where field research was carried out during this period. The list would be too long andcumbersome to include here. To all of you we give our sincere appreciation and deepest
gratitude.
The printing of this document was made possible through research funds provided toMichael A. Huffman (▉ 匁 C16570193) from the Ministry of Education, Science, Sports,
and Culture, Japan. My warmest appreciation goes to Fumie Naito and Reiko Nagata for
their kind help in scanning many of the papers included here within onto PDF files, which
made the final printing easier. Any mistakes or omissions are solely my responsibility.
Michael A. Huffman, Inuyama, Japan, January 2, 2005i

Every study has a beginning, and this is ours.
November 21, 1987, 4.50 am: The booming pant-hoots of a lone chimpanzee break the
pre-dawn stillness and wake me from a comfortable sleep. It is still nighttime in Mahale, but
the sun has already begun to rise over the Indian Ocean on the Tanzanian coast some 1000
km to the east, poised to begin its push across the expansive savannah woodlands of theinterior. The rays of early morning light will not reach Kansyana camp for at least anotherhour and a half. The Mahale Mountains are the last hurdle that the sun must overcome
before its welcome warm rays finally cover all of Tanzania.
Over the next two hours, the lone male urgently continues to call out in hope of getting a
reply from his scattered companions. Typical of Mahale chimpanzees during the rainyseason, they have begun to split into smaller groups in search of limited patches of food.
Now, at the beginning of the rainy season members of the Mahale M-group are not always
so easy to find.
At around 6:50, the male quietly passes through camp on his way north. There is lightenough now to see and I recognize him as Ntologi, M-group's alpha male since 1979.
At around 7:50, Mohamedi Seifu Kalunde, my friend and research assistant, arrives atcamp coming from Kasiha village. We waste no time and head north in the same direction
as Ntologi did. The first hour is spent moving north and just climbing up the steep slopes ofthe mountain to a high vantage point on the ridge where we are most likely to hearchimpanzees in the Mpila river valley-basin down below.
At 9:36, during our traverse of the forested G-ridge, we again hear the familiar series ofpant-hoots from a single male. Almost instantaneously thereafter, his pant-hoots are met by
a rally of replies in kind from a small party further up the river, followed by yet anothersmall group still further up the mountain. From their calls, we know that the chimpanzees
are split into small groups spread all across the forest, probably in search of fruits.
As quickly as they announce their presence, they are they swallowed up once again in thesilence within the thick dark green landscape.
Between 9:50 and 12:41, we find several individuals and begin to record their behavior,noting who is in the group, what they are feeding on, which females are sexually receptive
and so on.
This basic data has been accumulated on a daily basis since 1965 when the research firststarted.
Moshi Bunengwa, another of our team's research assistants, comes with the news that hehas seen Ntologi who is now traveling with two young males, Nsaba and Hito. Like
ourselves, Ntologi has successfully made contact with other group members.
Alone or in small groups, chimpanzees sit quietly in the canopy or hang one-handed frombranches, plucking out the red wax-membrane coated seeds of Lulumasia, a relative of thenutmeg, from the thick yellow husks clustered together like grapes. We first detect this small
party of chimpanzees only from the sound of discarded husks falling to the ground with alight thud. Chimpanzees on the ground are even harder to spot but they can be heard as theypull up stalks of Itungulu--African ginger or Iswe--elephant grass. Sometimes also
protruding out of the ground at the base of the stalks are the oblong red fruits of Itungulu, awelcome treat for chimpanzees and humans alike. Throughout the morning, individuals join
and split from the group we are watching.
At 12:41, Mohamedi and I hear a round of pant-grunt greetings and screams in the distance.
We intercept the group as they descend along the Mpila River. We find a group of elevenii

individuals including two adult males, Lukaja and Kasangazi, the then young adolescentmale Jilba, three adult females Chausiku, Wantendele and Wakilufya and their four
dependent offspring Chopin, Masudi, Maggy, and Ashula.
The three adult females, all immigrants from the now extinct Kajabala-group often travel
together when not sexually receptive. They have continued their friendships and help watch
over each other's young in Mimikire-group. They, no doubt, find solidarity amongst
themselves in a society where newcomer females are harassed by more dominant females incompetition for valuable food resources and possibly mates.
Wantendele, Wakilufya, and the adult male Lukaja are three of the target individuals in mystudy this season. Mohamedi and I begin our real work for the day at last. We start byfollowing Lukaja, the lead member of the group as he quickly moves down theriver-bed-obstacle-course of large slippery boulders in the Mpila riverbed. The rest of thechimpanzees follow closely behind. Lukaja and his two male companions are lost after only
16 minutes of contact. They leave the females behind as they double back and disappear into
thorny dense bush in the direction of pant-hoots from a larger group up on the ridge thesame place from where we first heard chimpanzees calling this morning.
We continue to follow the females as they quickly move into the forest then pause to listen to
the large group up on the ridge. They decide not to follow and begin to move at a leisurely
pace in the cool shade of the trees, in search of something to eat.
With extremely acute vision, Wantendele spots a cluster of ripe yellow Ilombo fruit high inthe canopy. She swiftly climbs up with Maggy clinging to her back. Her son, Masudi
continues to wrestle with Ashula and Chopin on the ground. Chausiku climbs up only ashort distance and rests. A hunch tells me that Chausiku does not seem to be feeling well.
She constructs a sturdy daybed by weaving live branches into a stable platform in the fork ofa large branch. She listlessly stares up into the canopy at her two-and-a-half-year-old son,as he precariously dangles from a branch by one hand. Masudi and Maggy tug at his feet in
an attempt to solicit play. If a young infant fell from such a height, it could easily be injured.
A mother is usually nearby to retrieve her infant and scold its older playmates, but today
Chausiku pays no attention. She appears weak and lacking in energy.
By 13:2,0 our initial group of eleven has dwindled down to only two adult females,
Wantendele and Chausiku, and their three offspring.
At 13:22, Wantendele moves south with Maggy and Masudi to feed on figs and more Ilombo.
Mohamedi and I decide to follow them. Wantendele doubles back and returns to the base of
the same tree where we left Chausiku 30 minutes earlier, as if she expects her to still be
there. She is and has not budged from her bed. Chopin continues playing alone higher up in
the branches.
At 14:02, Chausiku climbs down to the ground and meets Wantendele who has patiently
been waiting for her. They all start to move south. Chausiku's progress is painfully slow as
she frequently stops to rest. The others continue on ahead, pausing occasionally to feed on
the pith of Iswe and Itungulu, always remaining in sight of Chausiku.
At 14:13, Chausiku goes directly to and sits down in front of a shrub and pulls down several
new growth branches about the diameter of my little finger. She places them all on her lap
and removes the bark and leaves of the first branch to expose the succulent inner pith. She
then bites off small portions and chews on each for several seconds at a time. By doing this,she makes a conspicuous sucking sound as she extracts and swallows the juice, spitting out
most of the remaining fiber. This continues for 17 minutes, with short breaks as she
consumes the pith of each branch in the same manner.

iii

Not being familiar with this plant, I ask Mohamedi, as we continue to watch Chausiku, what
it is called in the local Tongwe vernacular. He replies, "Mjonso". Mohamedi has worked
here with the Kyoto University chimpanzee research project since it started in the early
1960s and is widely acknowledged by researchers and fellow Tongwe tribesman for his vast
knowledge of the flora and fauna in the area.
While Chausiku continues to feed, Wantendele sits nearby feeding on Iswe but shows nointerest in the Mjonso. Chopin does seem interested, however, and begs for a piece of pithhanging from his mother's mouth. He picks up a piece, which falls from her mouth and asection of the discarded leaf and bark from the ground. After putting these things in his
mouth for a brief second he quickly discards them and continues chewing on a piece of Iswe.
Not having seen chimpanzees eat this plant before, I next ask Mohamedi if chimpanzees
consume it often, and add that I have not seen them show any interest in it before today in myten months of observations here. He nods and replies that, indeed, it is not eaten frequently.
He adds that K-group chimpanzees also ate the pith, but that chimpanzees probably do not
eat too much because it is very bitter. Interested in the customs and material culture of his
people, I next ask him if the Tongwe use it for anything. He smiles and says matter of factly,"Yes, it is very strong Dawa (medicine)." "What kind of Dawa?", I ask in amazement. He
replies saying it is most commonly used for malaria-fever, stomachaches, and intestinal
parasites.
My mouth drops open as the details of our observations of Chausiku over the last hour anda half flash through my mind. I become consumed with an urgency to know just what theoutcome of Chausiku's behavior will be and I am filled with a rush of adrenaline. Without
speaking, we agree that we will follow and watch her closely. As time progresses, theseverity of her illness becomes more apparent. Her lower back appears painful because of
the careful and slow way she walks, sits down and climbs trees. She is also weak and has
little appetite. At one point we also note the unusually dark color of her urine as she relieves
herself off the side of her daybed.
Wantendele does not leave Chausiku for the remainder of the day as they slowly make their
way south to the northern foot of G-ridge.
At 15:01, Wantendele climbs up into a Lwago tree and feeds on its fruits. Chausiku slowly
and with obvious effort climbs up and makes a new daybed below Wantendele. Thirty
minutes later, Wantendele climbs down and sits on the ground. Chausiku raises her head,then slowly moves out of her bed and climbs partway down the tree, but stops and builds
another daybed. After a few minutes of hesitation, Wantendele climbs back up into the tree
and builds a daybed herself at 15:33; she remains there until Chausiku finally comes down
at 16:49 after another long rest.
Once again they set off south and Chausiku follows but quickly begins to lag behind theothers. After several stops, a drink of water and brief nibble on ginger pith, Chausiku
catches up with the others at 17:29. Wantendele, Maggy, and Masudi have already climbed
up into the trees for the night. Chopin sits waiting on the ground for his mother to arrive. In
a last struggle, Chausiku slowly climbs up into a tree and builds her last bed for the day at
17:44. Chopin climbs high up into the tree and explores on his own. We stay with them until
around 18:00 then head back to camp before it gets too dark.
On several occasions today we had the impression that Wantendele coordinated her
activities to keep an eye on Chausiku. The main group was occasionally within hearing
distance and probably less than a mile from us all afternoon. Nonetheless, Wantendele
made no attempt to leave her friend to join them.

by its Latin name as Vernonia amygdalina, but had neither associated its use with sickness
nor were they aware of the plant's medicinal value to the Tongwe. Their healthy skepticism
encourages me even more strongly to follow this through to the end.
The next morning between 7:00 and 8:00, a small group of three to four chimpanzees
pant-hoot on the mountain side Southeast of camp, as they find their first meal of the day.
Mohamedi and I head north.
By 9:00 we make contact with our first group of eleven chimpanzees. To our joy we findChausiku and Wantendele with them, only 60 meters from where they made their beds lastnight. Besides Chausiku, Wantendele and their offspring, included in the group is the higher
ranking adult male, Kalunde, the adolescent male Jilba, a recent immigrant, the young adultsexually receptive female Patty, and another adult female Gwekulo (born ca. 1962 inK-group).
Today Gwekulo seems to have the job of looking after Chopin. This is not the first time for
Gwekulo to watch over Chausiku's infants in times of sickness. Such associations are known
as 'baby-sitter relationships' in which young adult females travel together with a particularmother-infant pair and act as a playmate for the infant and a travel companion for themother. When these young females are sexually receptive, however, they usually travel
within groups, which also contain males. Gwekulo never had the luck to have children ofher own. Thus, even nowadays she very often moves around with other females and theiroffspring.
While Chausiku is lying down off to the side of the group, Gwekulo sits nearby Chopin,
watching over him as he wrestles with Ashula and Masudi. Occasionally, Patty solicits
Chopin to copulate. At 9:59, the group moves south. Chopin follows after Patty. Chausiku
slowly follows behind the group for less than a minute before she separates from Chopin
and the rest for the next 38 minutes. She still appears quite weak, frequently stopping to sit
and rest.
At 10:42 she is seen to eat for the first time today. Chopin is now traveling with his motheragain and between 11:03 and 11:31 they slowly follow behind the group feeding on small
amounts of Kasolyo figs and Lulumasia.
Chausiku catches up with the rest of the group and they all stop for their habitual midday
nap.
At 12:49, after a long 72 minute sleep, everyone awakens in a commotion of pant-hoots andscreams as if a fire station alarm had just gone off in the middle of the night and is up and on
the move. Mohamedi and I are caught off guard. As they disappear into the tangle of thornyvines, we crawl on hands and knees, sometimes squirming on our bellies, to keep up withthem.
At 13:06, we succeed in getting through the worst of it to find the group nonchalantly resting
again. Chausiku, Chopin, and Gwekulo, however, immediately get up and leave the rest of
the group behind as they head southwest.
Following behind at a half running pace just to keep up with Chausiku, it becomes clear tous that her condition is improving. She pauses occasionally, this time looking behind her, tosee if the others are keeping up. Chausiku seems irritable and distracted, slapping the
ground and grunting when Chopin refuses to follow close behind. Gwekulo stays behind to
wait for Chopin to catch up while Chausiku continues to move on ahead.

Back at camp, our story is met with both excitement and disbelief by some of the senior
members of our research team. They had, of course, seen chimpanzees use this plant, known

Finally at 13:51, they all end up in an open semi-swamp grassland area filled with three tofour meter high elephant grass. Chausiku continues to eat steadily, consuming the succulent
pith of elephant grass and ginger, the small semi-sweet red figs of Kankolonkombe and theivv

tangy baseball-sized yellow fruits of Ilombo. We terminate our observations at 16:05, when
we can no longer keep up with her.
What Mohamedi and I witnessed over these days was to be the beginning of a study that hascontinued for the last decade. Since this time we have observed other sick individuals
consume the bitter pith of Mjonso and we have found still other plants like Aspilia that may
be used in other ways as medicine by these chimpanzees.
The production of toxins, drugs and other compounds, called secondary metabolites by thechemists who study them, is considered to be an evolutionary adaptation to help plants fightoff predation from insects and herbivores. These compounds therefore greatly influence
what plants animals can select as food and animal ecologists have focused a great deal of
their research on understanding how animals can cope with such compounds.
Mohamedi, during my years of field work at Mahale, has been a very valuable friend andcollaborator. He was taught herbal medicine by his mother and grandfather--babu
Kalunde; both were traditional healers of his community. Had it not been for the knowledge
and investment made by Mohamedi’s family in him, I am sure that the story I have just told
you would never had happened.
Mohamedi once told me that when he was a boy, babu Kalunde told him a story about how
he acquired an important new medicine for humans by watching the behavior of a sickanimal.
The story goes that babu Kalunde had taken in a young porcupine after its mother wascaught and killed in a snare. Shortly afterwards, the young porcupine became sick, andsuffered from diarrhea and lethargy. It wandered off from the village and babu Kalunde
followed him. The porcupine dug up the root of a plant the Tongwe call Mulengelele. Notingthat the baby porcupine recovered from its illness, babu Kalunde decided to collect some ofthese roots and try them out on people in his village, who, too, had fallen ill. After thatexperiment Mulengelele became an important plant for the treatment of parasites amongst
the Tongwe.

Mohamedi, Huffman and Luhembe
Ishamili at Kansyana camp (December
1995).

Over the history of humankind, people developed elaborate health-care systems based on
experimentation and observations of nature. We in the ‘developed world’ have much tolearn from the people of traditionally living cultures as well as from our primate cousins
who continue to live closely with nature.
What Mohamedi and I witnessed over those two days in 1987 was the beginning of a very
productive study that has continued to this day and promises to go on for a long time tocome. I have continued to expand the scope of this research and I am now collaboratingwith colleagues working at a number of other field sites throughout Africa and Asia. The
laboratory side of this research now includes collaborations with pharmacologists,
parasitologists, veterinarians, physiologist, botanists and others in countries around theglobe such as Japan, Tanzania, Uganda, United States, Canada, United Kingdom, Australia,
France, Holland, Denmark, Germany, Saudi Arabia, and the Czech Republic. Our goal is
not only to better understand how animals deal with illness, but also to search for new
medicines and ecologically sound strategies for the treatment of major parasite diseases
like malaria, schistosomiasis, lieshmania, and others.
Vernonia amygdalina has proven to be a very important plant in this endeavor. It is our goalthat the research will lead to the development of new strategies for treating human andlivestock parasite disease based on adaptive behaviors which have proven the test of time inthe struggle for survival between host and parasite. (An excerpt taken from Hofer,
Huffman & Gunter 2000 MAHALE- A photographic encounter with chimpanzees. With a
Forward by Jane Goodall, Sterling Press, NY pp. 110-117.

vi

Meeting with our colleagues in the Department of Chemistry, University of Dar es Salaam.
Left to right: Koichi Koshimizu, Cosam Joseph, Huffman, Mayunga H.H. Nkunya, Hajime
Ohigashi, Hideo Hayashi (October 1999),

vii

Contents
1.

Huffman, M. A. & Seifu, M. (1989). Observations on the illness and consumption of a
possibly medicinal plant Vernonia amygdalina by a wild chimpanzee in the Mahale
Mountains, Tanzania. Primates 30(1): 51-63.
….…..1

2.

Ohigashi, H., Takagaki, T., Koshimizu, K. Nishida, T., Huffman, M. A., Takasaki, H.,
Jato, J. & Muanza, D.N. (1991). Biological activities of plant extracts from tropical
Africa. African Study Monographs 12(4): 201-210.
….......14

3.

Ohigashi, H., Jisaka, M., Takagaki, T., Nozaki, H., Tada, T., Huffman, M. A., Nishida,T., Kaji, M. & Koshimizu. K. (1991). Bitter principle and a related steroid glucoside of
Vernonia amygdalina, a possible medicinal plant for wild chimpanzees. Agricultural
and Biological Chemistry. 55(4): 1201-1203.
.......... 24

4.

Ohigashi, H., Jisaka, M., Takagaki, T., Koshimizu, K., Huffman, M. A., Nishida, T.,
Takasaki, H. & Kaji, M. (1991). Possible medicinal plants used by wild chimpanzees,
and their physiologically active compounds. Journal of African Studies 39: 15-27. (InJapanese) 啾些ⅺ↹ ↗ ↹ ⅱҚ ℚ 善嗩㏬双℔ 些噃┣二亍収, Ҵ 1⁄8 ↛ ↰ Ⅴ➂◧ ҵ 39:
15-27.
...........27

5.

Jisaka, M., Kawanaka, M., Sugiyama, H., Takegawa, K., Huffman, M. A., Ohigashi, H.
& Koshimizu, K. (1992). Antischistosomal activities of sesquiterpene lactones andsteroid glucosides from V. amygdalina, possibly used by wild chimpanzees against
parasite-related disease. Bioscience, Biotechnology and Biochemistry 56(5): 845-846.
..........40

6.

Jisaka, M., Ohigashi, H., Takagaki, T., Nozaki, H., Tada, T., Hirota, M., Irie, R.,
Huffman, M.A., Nishida, T., Kaji, M. & Koshimizu, K. (1992). Bitter steroid
glucosides, Vernoniosides A1, A2 and A3 and related B1 from a possible medicinal
plant Vernonia amygdalina, used by wild chimpanzees. Tetrahedron 48: 625-632.
.........427.

Huffman, M. A., Gotoh, S., Izutsu, D., Koshimizu, K. & Kalunde, M. S. (1993).
Further observations on the use of the medicinal plant, Vernonia amygdalina (Del) by awild chimpanzee, its possible effect on parasite load, and its phytochemistry. African
Study Monographs 14(4): 227-240.
.........50

8.

Koshimizu, K., Ohigashi, H., Huffman, M. A., Nishida, T. & Takasaki, H. (1993).
Physiological activities and the active constituents of potentially medicinal plants usedby wild chimpanzees of the Mahale Mountains, Tanzania. International Journal ofPrimatology 14(2): 345-356.
........64

9.

Jisaka, M., Ohigashi, H., Takegawa, K., Huffman, M. A. & Koshimizu, K. (1993).
Antitumoral and antimicrobial activities of bitter sesquiterpene lactones of Vernonia
amygdalina, a possible medicinal plant used by wild chimpanzee. Bioscience,
Biotechnology, Biochemistry 57(5): 833-834.
........76

This volume is dedicated to the memory of Chausiku (ca. 1958 1990), my first mentor in chimpanzee self-medication, and ateacher to many of us who have had the privileged of sharing
time with Mahale chimpanzees. (photo by M.A. Huffman, Nov.
1987).

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ix

10. Jisaka, M., Ohigashi, H., Takegawa, H., Hirota, M., Irie, R., Huffman, M. A. &Koshimizu. K (1993). Steroid glucosides from Vernonia amygdalina, a possible
chimpanzee medicinal plant. Phytochemistry 34(2): 409-413.
……….78

11. Huffman, M. A. (1993). An investigation of the use of medicinal plants by wild

chimpanzees. Current status and future prospects. Primate Research 9:179-187. (inJapanese with English Abstract) 啾些ⅺ↹ ↗ ↹ ⅱҚ ℚ 善佻嘽嗩➂◧ Ѽ 亍⒔℔ 充呻ѷҴ 囏僄嚼➂◧ ҵ 9Ѽ 179-187......... .83

12. Gasquet, M., Huffman, M. A., & Wrangham, R. W. (1994). Les plantes medicinales
utilises par les chimpanzes sauvages. Metissages en Sante Animale (Eds.) K. Kasonia& M. Ansay, Namur: Presses Universitaires de Namur (Belgium), pp. 289-297. (inFrench)…….. 92

13. Ohigashi, H., Huffman, M. A., Izutsu, D., Koshimizu, K., Kawanaka, M., Sugiyama,

H., Kirby, G. C., Warhurst, D. C., Allen, D., Wright, C. W., Phillipson, J. D.,
Timmon-David, P., Delmas, F., Elias, R., Balansard, G. (1994). Toward the chemical
ecology of medicinal plant use in chimpanzees: The case of Vernonia amygdalina, aplant used by wild chimpanzees possibly for parasite-related diseases. Journal ofChemical Ecology 20(3): 541-553.
............ 102

20. Ohigashi, H., Huffman, M.A., Koshimizu, K. (1996). In the search for potential
medicines from the tropics: interfacing chimpanzee behavior and African
Ethnomedicine. In: Namba, T. & Kadota, S. (Eds.) Proceedings of the 5thInternational Symposium on Traditional Medicine in Toyama (1996). Hosted by the
Research Institute for Sino-Japanese Medicine Research & Toyama Medical andPharmaceutical University, Chuo Publishers, Toyama, pp. 165-174.
...........196

21. Huffman, M. A., Koshimizu, K. & Ohigashi, H.

(1996). Ethnobotany andzoopharmacognosy of Vernonia amygdalina, a medicinal plant used by humans andchimpanzees. In: PDS Caligari and DJN Hind (Eds.). Compositae: Biology &Utilization Vol 2., Kew: The Royal Botanical Gardens, pp. 351-360.
...... 207

22. Ohigashi, H., Huffman, M. A., & Koshimizu, K. (1996). Chemical ecology of the

medicinal use of Vernonia amygdalina by chimpanzees. Journal of African Studies 48:
51-62. (in Japanese) (in Japanese) 啾些ⅺ↹ ↗ ↹ ⅱҚ ℚ 善嗩㏬双嘽嗩℗ ╱ ℅ ℷ ⒅┓些俗┓儴➂◧ : Vernonia amygdalina 4⁄5 囁℗ , Ҵ 1⁄8 ↛ ↰ Ⅴ➂◧ ҵ 48: 51-62.
...........218

23. Huffman, M. A., Gotoh, S. Turner, L. A., Hamai, M. & Yoshida, K. (1997). Seasonal
trends in intestinal nematode infection and medicinal plant use among chimpanzees in
the Mahale Mountains National Park, Tanzania. Primates 38(2): 111-125.
...........230

14. Koshimizu, K., Ohigashi, H. & Huffman, M. A. (1994). Use of Vernonia amygdalina

24. Page, J.E., Huffman, M. A., Smith, V. & Towers, G. H. N. (1997). Chemical basis for

15. Huffman, M. A. & Wrangham, R. W. (1994). Diversity of medicinal plant use by

25. Huffman, M. A. (1997). Current evidence for self-medication in primates: aby wild chimpanzees; Possible roles of its bitter and related constituents. Physiology
and Behavior 5(6): 1209-1216.
........... 115
chimpanzees in the wild. In Chimpanzee Cultures. (Eds.) R.W. Wrangham, W.C.
McGrew, F.B. de Waal & P.G. Heltne, Harvard Univ. Press, Mass. pp. 129-148.
...........123

16. Huffman, M. A. (1994). The C.H.I.M.P.P. Group: A multi-disciplinary investigation

Aspilia leaf-swallowing by chimpanzees: a re-analysis. Journal of Chemical Ecology
23(9): 2211-2226.
...........245
multidisciplinary perspective. Yearbook of Physical Anthropology 40: 171-200.
...........261

26. Huffman, M. A. (1997). Practical applications from the study of great ape
self-medication and conservation related issues. Pan African News 4(2): 15-16.
...........291

into the use of medicinal plants by chimpanzees. Pan Africa News 1(1): 3-5.

...........143

17. Huffman, M. A. (1995). La pharmacope des chimpanzes. La Recherche October Issue,
pp. 66-71. (in French)..........147

18. Mahaney, C. W., Hancock, R. G. V., Aufreiter, S. & Huffman, M. A. (1996).

Geochemistry and clay mineralogy of termite mound soil and the role of geophagy inchimpanzees of the Mahale Mountains, Tanzania. Primates 37(2): 121-134.
...........153

19. Huffman, M. A., Page, J. E., Sukhdeo, M. V. K., Gotoh, S., Kalunde, M. S.,
Chandrasiri, T. & Towers, G. H. N. (1996). Leaf-swallowing by chimpanzees, abehavioral adaptation for the control of strongyle nematode infections. International
Journal of Primatolology 17(4): 475-503.
...........167x

27. Huffman, M. A., Ohigashi H., Kawanaka, M., Page, J. E., Kirby G. C., Gasquet, M.,
Murakami, A., & Koshimizu, K. (1998). African great ape self-medication: A newparadigm for treating parasite disease with natural medicines? In: Towards Natural
Medicine Research in the 21st Century, (Ed.) Y. Ebizuka, Amsterdam: Elsevier
Science B.V., pp. 113-123.
..........293

28. Huffman, M. A., Riad, E., Ohigashi, H. & Nansen, P. (1998). Self-medication in the
Great Apes: A multidisciplinary study of diet, behavior and health. Primatologie 1:
179-204. (French)..........304

29. Huffman, M. A. (1998). Control of nematode infections by African great apes: a newparadigm for treating parasite infection with natural medicines? American Associationof Veterinary Parasitologists. 20(2): 3-7.
.........330xi

30. Gasser R. B., Woods W. G., Huffman M. A., Blotkamp J. & Polderman, A.M. (1999).
Molecular separation of Oesophagostomum stephanostomum and Oesophagostomum
bifurcum (Nematoda: Strongyloidea) from non-human primates. International Journal
for Parasitology 29: 1087-1091.
...........336

31. Mahaney, W. C., Zippin, J., Milner M. A., Sanmugadas, K., Hancock, R. G. V.,

Aufreiter, S., Campbell, S., Huffman, M. A. & Wink, M. (1999). Chemistry,mineralogy and microbiology of termite mound soil eaten by chimpanzees of theMahale Mountains, Western Tanzania. Journal of Tropical Ecology 15: 565-588.
...........341

32. Alados C. L. & Huffman, M. A. (2000). Fractal long range correlations in behaviouralsequences of wild chimpanzees: a non-invasive analytical tool for the evaluation ofhealth. Ethology 106: 105-116.
..........365

33. Huffman, M. A., Koshimizu, K. & Ohigashi, H. (2000). The medicinal diet of primates.

Kyoto University Press, Kyoto (Japanese), In Primate Ecology- the dynamics ofenvironment and behavior. (Ed.) Y. Sugiyama, Kyoto: Kyoto University Press, pp.
85-108.Ҩ in Japaneseҩ Ⅾ↱ ℚ 善伱嚅噃, Ҵ 囏僄嚼些俗┓: ╝ ☢℔ に冽ℚ ⅹ5⁄8 ←↥ⅳ↦ ҵ , 乣㈯ぃ╶ 吂, ☏八俳┓┓㍙㍘勪Ⓓ, ☏八, pp. 85-108.
........377

34. Huffman, M. A. (2000). Forest Pharmacy. Healthy Options March Issue: 10-14

New evidence for leaf swallowing and Oesophagostomum infections in bonobos (Panpaniscus),International Journal of Primatology� 23 (5):1053-1062.
.... .493

41. Cousins, D. & Huffman, M. A. (2002). Medicinal properties in the diet of gorillas- an
ethnopharmacological evaluation. African Study Monographs� 23:65-89.
..........503

42. Huffman, M. A., Ohigashi, H. & Koshimizu, K. (2002). Multidisciplinary research on

chimpanzees self-medication. In The Mahale Chimpanzees- Thirty seven years ofPanthropology (Eds.) T. Nishida & K. Kawanaka, S. Uehara, Kyoto: Kyoto University
Press, pp. 261 -287. ㋔➩㋈嚊に冽ℚ ┓ペ儴➂◧ ,Ҵ ↤ ↕ ↲ ℚ ⅺ↹ ↗ ↹ ⅱҚ ↗ ↹ⅲ↳ ↣ ↳ ⅰҚ ℚ ㈬㋠削ҵ ѷ 亡兒嘽儆ѷ ㏏➒メートル偉ѷ 仮偲❢初吂傒ѷ ☏八俳┓┓㍙㍘勪Ⓓ, ☏八, pp. 261-287.
……..528

43. Huffman, M.A. (2002) Animal origins of herbal medicine. (Origines animales de la

medicine par les plantes) In. Des sources du savoir aux medicaments du futur- Fromthe sources of knowledge to the medicines of the future. (Eds.) J. Fleurentin, J-M. Pelt,
G. Mazars, Paris: IRD Editions, (English) pp. 31-42, (French) pp. 43-54.
..........555

44. Carrai, V., Borgognini-Tarli, S. M., Huffman, M.A., Bardi, M. (2003). Increase in........401

35. Huffman, M. A. & Caton J. M. (2001). Self-induced increase of gut motility and thecontrol of parasitic infections in wild chimpanzees.
Primatology 22(3) 329-346.

40. Dupain, J., van Elsaker, L., Nell, C., Garcia, P., Ponce, F. & Huffman, M. A. (2002).

International Journal of……..406

36. Wakibara, J. V., Huffman, M. A., Wink, M, Reich, S., Aufreiter, S., Hancock, R. G. V.,

Sodhi, R., Mahaney, W. C. & Russell, S. (2001). Adaptive Significance of Geophagy
for Japanese Macaques (Macaca fuscata) at Arashiyama, Japan. International Journalof Primatology 22 (3): 495-520.
..........424

37. Aufreiter, S., Mahaney, W. C., Milner, M. W., Huffman, M. A., Hancock, R. G. V.,
Wink, M., Reich, M. & Rana, S. (2001). Mineralogical and chemical interactions ofsoils eaten by chimpanzees of the Mahale Mountains and Gombe Stream National
Parks, Tanzania. Journal of Chemical Ecology 27(2): 285-311.
..........450

38. Ketch, L. A., Malloch, D., Mahaney, W. C. & Huffman, M. A. (2001). Comparative

microbial analysis and clay mineralogy of soils eaten by chimpanzees (Pan troglodytes
schweinfurthii) in the Mahale Mountains National Park, Tanzania. Soil Biology andBiochemistry 33(2): 199-203.
..........477

tannin consumption by sifaka (Propithecus verreauxi verreauxi) females during thebirth season: a case for self-medication in prosimians? Primates 44(1): 61-66.
..........579

45. Huffman, M. A. & Hirata, S. (2003). Biological and Ecological Foundations of Primate

Behavioral Traditions. In. The Biology of Tradions. (Eds.) D. M. Fragaszy & S. Perry,
Cambridge: University of Cambridge Press, pp. 267-296.
.........585

46. Huffman, M.A. (2003). Animal self-medication and ethnomedicine: exploration andexploitation of the medicinal properties of plants. Proceeding of the Nutritional Society
62: 371-381.
........615

47. Huffman, M.A. & Hirata, S. (2004). An experimental study of leaf swallowing in

captive chimpanzees- insights into the origin of a self-medicative behavior and the roleof social learning. Primates 45(2): 113-118.
……..626

48. Kaur, T., Huffman, M.A. (2004). Descriptive urological record of chimpanzees (Pantroglodytes schweinfurthii) in the wild and limitations associated with using
multi-reagent dipstick test strips. Journal of Medical Primatology� 33(4): 187-196.
........632

39. Huffman, M. A. (2001). Self-medicative behavior in the African Great Apes: An

evolutionary perspective into the origins of human traditional medicine. BioScience
51(8): 651-661.
..........482

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Additional publications and manuscripts in press / under review not printed infull in this document
Hofer, A., Huffman, M. A, & Ziesler, G. (1998). Mahale - Begegnung mit Schimpansen.
PAN Edition im Verlag Navalon, Fussen
Hofer, A., Huffman, M. A, & Ziesler, G. (2000) Mahale; a photographic encounter withchimpanzees. Sterling Publishing, New York.
Huffman, M.A. (2005 in press) Animal self-medicative behavior. In. The Encyclopedia of
Animal Behavior. Greenwood Publishers, Westport CT
Huffman, M.A., Cousins, D. (2005 in press). Potential medicative value in the diet ofgorillas. In GRASP World Great Ape Atlas.
Huffman, M.A., and Vitazkova, S.. (2005 in press). Primates, plants, and parasites: theevolution of animal self-medication and ethnomedicine. In Encyclopedia of Life Support
Systems. (eds.) N. Etkin & E. Elisabetsky UNESCO-Eolss Publishers Co. Ltd., Paris.
Krief, S., M. A. Huffman, T. Sévenet, J. Guillot, C. Bories, C. M. Hladik and R. W.
Wrangham (2005 in press) Non-invasive monitoring of the health condition of wild
chimpanzees (Pan troglodytes schweinfurthii) in the Kibale National Park, Uganda.
International Journal of Primatology 26:
Mnason, T., Reynold, V., Huffman, M.A., Pebsworth, P., Mahaney, W.C., Milner, M.,
Waddell, A., Dirszowsky, R., Hancock, G.V. (accepted). Geophagy in chimpanzees (Pantroglodytes schweinfurthii) of the Budongo forest reserve, Uganda. In: Primate of Western
Uganda, J. Paterson, V. Reynolds, H. Grant, N., Newton-Fisher, N. E. (eds.), pp. X – X.
Pebsworth, P., Krief, S., Huffman, M.A. (accepted) The Role of Diet in Self-Medication
Among Chimpanzees in the Sonso and Kanyawara Communitites, Uganda. in chimpanzees(Pan troglodytes schweinfurthii) of the Budongo forest reserve, Uganda. In: Primate ofWestern Uganda, J. Paterson, V. Reynolds, H. Grant, N., Newton-Fisher, N. E. (eds.), pp. X– X.
Krief, S., Huffman, M.A., Sévenet, T., Hladik, C-M, Grellier, P., Philippe Loiseau, M.,
Wrangham, R.W. (under review) Bioactive Properties of Plants ingested by Chimpanzees(Pan troglodytes schweinfurthii) in the Kibale National Park, Uganda. American Journal ofPrimatology
Petrezelková, K. J., Hasegawa, H., Moscovice L, R., Kaur , T., Issa M., and Huffman, M. A.
(under review). New records of parasitic nematodes for chimpanzees found from theintroduced population on Rubondo Island, Tanzania. International Journal of Primatology
Huffman, M. A., (under review) Primate Self-Medication. In: Primates in Perspective,
Christina Campbell, Agustin Fuentes, Katherine MacKinnon, Melissa Panger, Simon
Bearder (eds.), Press

Public Education
Examples of the research introduced in academic textbooks, technical literature and popularscientific literature:
The Plant-Book. A portable dictionary of the vascular plants. 2nd Edition. (1997). D. J.
Mabberley, Cambridge University Press, Cambridge. Vernonia amygdalina description
includes use of chimpanzees when sick and bioactivities properties of pith. (p. 744).
Parasitology & Vector Biology 2nd Edition (2000). William C. Marquardt, Richard S.
Demaree, Robert B. Grieve, Harcourt Academic Press, San Diego. “Self-treatment forparasites” (text box p. 390).
The Natural History of Medicinal Plants (2000). Judith Sumner, Timber Press, Portland.
“Chimpanzees and self-medication” (pp. 149-150).
Natures Medicine Plants that Heal (2000) Joel L. Sherdlow, National Geographic Society,
Washington, D.C. “Vernonia amygdalina, includes description of medicinal uses forhumans and chimpanzees. (p. 388). On animal self-medication (pp. 176-177, 180).
Parasite Rex. Inside the world of nature’s most dangerous creatures (2000). Carl Zimmer,
The Free Press, New York (pp. 201-202).
Medicine Quest. In search of Nature’s Healing Secrets (2000). Mark J. Plotkin, Viking
Penguin Press, New York, “Plants of the Apes” (pp. 163-165, 167-169).
Wild Health. How animals keep themselves well and what we can learn from them (2002)
Cindy Engel, Houghton Mifflin Company, Boston, “chimpanzees” (pp. 46, 200), parasites(pp. 139, 140, 142-143, 146), “self-medication, determination of” (p. 37��� � �冽双� � � ㋔
伝��呄� � 啾些� 偎�� �� �� �� ��� �� ⅰ� �� � �� ⅱ�� �� 兒仚㊊喆�� � 妝��兆�
Primate Behavioral Ecology 1st & 2nd Editions (2000, 2003). Karen Strier, Allyn and Bacon,
Boston. “”Box 6.1 Forest Pharmacy” (pp. 180-181).
Parasites and the Behavior of Animals (2002). Janice Moore, Oxford Series in Ecology and
Evolution, Oxford University Press, Oxford. “Self-medication” (pp. 135-137).
Modern Biology (2005 Edition) Holt, Rinehart and Winston, Austin. Science in Action
Feature “Do animals self-medicate? (p. 598).
Representative account of research featured in international film & radio
documentaries“Wisdom of the Wild”(2000), Argo Productions Washington D.C. for Public Broadcasting
System (USA). An national and later internationally televised documentary with special
feature of chimpanzee self-medication research project at Mahale.
“Nature's Secrets” (2002), Discovery Channel, Toronto. An internationally televised
documentary with special feature of chimpanzee self-medication research at Mahale .
“Dr. Animal” (2003), Tele Image Paris for Discovery Channel, Tonronto. Aninternationally televised documentary, with special feature of chimpanzee self-medication
research at Mahale.

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“Animal Medicine” St. Thomas, Prod., Aix en Provence, France for National Geographic
Television (2004) Documentary on self-medicative behavior in the animal kingdom.
“Magic, murder, and medicine”, BBC Radio 4 (2001), Interview for 2 part series on
medicinal plants.
“Animal self-medication”, BBC Radio 4 (2003), A special program on animal
self-medication.

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