Insec applica ions o open wounds
by chimpanzees in he wild:
i s insigh s om Eas A ican
chimpanzees
Kayla Kol 1, Daniela Acos a Fló ez2, Alessand a Masca o1 & Simone Pika1
Medica i e beha iou s a e widesp ead among animals, and chimpanzees in he wild may exhibi a
newly iden i ied o m in ol ing he applica ion o insec s o open wounds. To da e, insec applica ions
o wounds ha e only been epo ed in a single communi y o Cen al chimpanzees (Pan oglody es
oglody es). Thus, we epo obse a ions o simila beha iou s in Eas e n chimpanzees (P. .
schwein u hii) o he Ngogo chimpanzee popula ion, in Kibale Na ional Pa k, Uganda. Be ween
No embe 2021 and July 2022, we obse ed six indi iduals ( h ee males, h ee emales) applying
lying insec s o hei own wounds (N = 5) and, in one ins ance, o he wound o a conspeci ic. These
obse a ions demons a e a gene ally consis en sequen ial pa e n o insec applica ions in bo h
Eas e n and Cen al chimpanzees. Al hough he ex en and po en ial medicinal unc ion o his
beha iou emain unclea , we p opose h ee hypo heses o in o m u u e esea ch, ocusing on
insec selec i i y, social ansmission, and p osociali y. In conclusion, he indings sugges ha insec
applica ions a e mo e widesp ead han cu en ly documen ed, and p o ide a basis o in es iga ing
hei acquisi ion, social dynamics, and po en ial ele ance o he e olu ion o human medicinal
beha iou s.
Keywo ds Animal medica ion, Sel -medica ion, O he - ega ding ca e, Pan oglody es schwein u hii,
P osocial beha iou , En omo he apy
T adi ional medicinal beha iou s a e a cul u al cha ac e is ic1 ha human socie ies ha e assembled o e
housands o yea s2–4. These beha iou s ha e been passed on h ough o al ansmission, obse a ion, and di ec
copying (i.e., social lea ning)5, wi h adi ional medicine con inuing o play a i al ole in global heal hca e6,7
(bu see also8,9. Unde s anding he o igins o hese medicinal beha iou s – such as he de elopmen o use o
he bal medicines, oin men s, en omo he apy ( he medicinal use o insec s3, and mode n pha maceu icals –
equi es examining how knowledge o medicinal compounds may ha e eme ged, been ansmi ed, and e ined
o e ime.
E idence sugges s ha medicinal plan s and ungi we e used by ou ex inc hominin ances o s, he
Neande hals (Homo neande halensis) as ea ly as 50,000 yea s ago, wi h some o hese plan s s ill in use
oday4,10. Fo ins ance, Ge man chamomile (Ma ica ia chamomilla) shown o ha e li le nu i ional alue10, and
oday used o di e en pu poses including an i-in lamma o y, an ispasmodic, and seda i e e ec s, as well as he
ea men o gas op o ec i e dis u bances11. Simila ly, en omo he apy3,12,13 has been used in some cul u es o
cen u ies. Fo ins ance, in China, app oxima ely 300 insec species ac oss 70 di e en gene a ha e been iden i ied
as ha ing medicinal p ope ies and a e included in adi ional Chinese medicine14. Simila ly, in Bu kina Faso,
adi ional heale s use insec species om six di e en o de s o a a ie y o medicinal pu poses15. The insec s
a e used o ea a wide ange o ailmen s, including diges i e and skin diso de s3, and hei documen ed e ec s
en ail an i-in lamma o y, an imic obial, an i i al, immunological, analgesic, an ibac e ial, diu e ic, anaes he ic,
and an i heuma ic e ec s3,12.
This aises a cen al ques ion: how did humans i s ecognise and ha ness such medicinal p ope ies, and how
migh his ha e e ol ed? One explana ion, sugges ed by schola s, is ha h oughou human his o y, people ha e
obse ed nonhuman animals (he ea e animals) ea ing hei illnesses using plan seconda y compounds, non-
nu i ional subs ances, o bioac i e subs ances16–18. These obse a ions ha e con ibu ed o he iden i ica ion
1Compa a i e BioCogni ion, Ins i u e o Cogni i e Science, Osnab ück Uni e si y, Osnab ück, Ge many.
2Depa men o An h opology, Facul y o A s and Sciences, Yale uni e si y, New Ha en, Uni ed S a es o Ame ica.
email: kk[email p o ec ed]
OPEN
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o plan -based ea men s wi h po en ial bene i s o humans19. This explana ion aises a u he ques ion: how
did such beha iou s eme ge in animals hemsel es? A g owing body o esea ch ac oss he animal kingdom –
om in e eb a es o p ima es20–30 – has shown ha a ious animals engage in medica i e beha iou s. These
include he use o na u al subs ances such as plan ma e 31–33, clay29, o millipedes (Sechellep us spp.)34 o o
p ophylac ic o he apeu ic pu poses, o en in ol ing an ipa asi ic e ec s (see also35.
Mos exis ing esea ch has ocused on cases o sel -medica ion in ol ing plan ma e ials, wi h only limi ed
examples o animal-de i ed subs ances35. Fu he mo e, un il ecen ly, he use o medicinal subs ances o ea
o he s was hough o be a e in animals, howe e , eme ging e idence sugges s i may be mo e widesp ead
han p e iously assumed35–37. These indings con ibu e o he ongoing deba e abou he exis ence o p osocial
beha iou s in nonhuman p ima es38,39, de ined as ac ions in ended o bene i ano he and, in humans, p ima ily
d i en by empa hic conce ns o o he s40,41. Hence, he social con ex o wound ending may ep esen a c ucial
candida e o enhancing ou unde s anding o p osocial and he apeu ic beha iou s in animals and he e olu ion
o human medicinal ca e.
One compelling example o such beha iou comes om ecen obse a ions on Cen al chimpanzees (Pan
oglody es oglody es) om he Loango Na ional Pa k, Gabon. These obse a ions documen ed a po en ially
signi ican example o bo h sel - and o he -di ec ed medica ion in ol ing he applica ion o insec s o open
wounds – no only hei own bu also hose o ela ed and un ela ed g oup membe s37. O e a pe iod o 15
mon hs, a o al o 19 sel -di ec ed e en s ( i e adul males, one adul emale, one ju enile emale) and h ee
o he -di ec ed e en s (one emale, wo males; one kin-di ec ed and wo non-kin-di ec ed e en s) we e obse ed.
These e en s ollowed a consis en beha iou al sequence:
• A lying insec was caugh .
• I was immobilised by being placed and/o squeezed be ween he lips.
• The insec was applied o he exposed su ace o he wound wi h he inge s and mo ed ac oss i using he
inge s o lips.
• I was emo ed om he wound using ei he he mou h o inge s and disca ded.
These obse a ions expand ou exis ing knowledge o animal medica ion o include allo-medica ion o open
wounds in ol ing he use o animal ma e 37. In addi ion, since documen ed e en s o insec applica ion in
chimpanzees in he wild ha e only ecen ly been obse ed, ques ions a ise ega ding he beha iou ’s uniqueness,
dis ibu ion, con ex -speci ici y, and links o ecological and sociodemog aphic ac o s. Unde s anding hese
aspec s helps de e mine whe he he beha iou e lec s an in insic esponse, a communi y-speci ic inno a ion,
o pa o a b oade pa e n, wi h implica ions o chimpanzee sel -medica ion, social ansmission, and p osocial
beha iou . He e, we p o ide i s insigh s in o insec applica ion in ano he chimpanzee subspecies – Eas e n
chimpanzees (Pan oglody es schwein u hii) – om he Ngogo popula ion, Kibale Na ional Pa k, Uganda.
We epo h ee e en s: wo e en s o sel -di ec ed applica ion, and one o o he -di ec ed applica ion. These
obse a ions o e compa a i e insigh in o he dis ibu ion and signi icance o insec -based wound ea men in
chimpanzees and i s ele ance o he e olu ion o p osocial and heal h- ela ed beha iou s.
Resul s
In o al, we obse ed i e e en s in ol ing insec - ela ed beha iou wi h wounds. These e en s a ied in e ms
o indi idual iden i y, sex, and age class o he applica o , wound loca ion, and pee ing occu ence (Table1). We
u he epo on h ee e en s due o he a ailabili y o ideo oo age and isibili y, en ailing wo sel -di ec ed
applica ions, and one o he -di ec ed applica ion.
E en 1 (see Video S1). An adolescen emale chimpanzee, F icka, sel -applied a lying insec o
an open wound on he inne igh high
This e en in ol ed an adolescen emale, F icka, who was obse ed applying an immobilised insec o an open
wound on he inne igh high (app oxima ely < 5cm in diame e ; Fig.1; Video S1, 00:16 − 01:42). She p essed
Da e ID Sex Age class Wound loca ion Applica o Pee ing Video oo age Example e en s
Wes e n communi y
23/11/21 Damien M Adolescen Le leg Sel Yes Yes 2
23/11/21 Damien M Adolescen Le leg E a James Yes Yes 3
Cen al Communi y
28/11/21 Clay on M Adolescen Le leg Sel No No
3/12/21 Ab ams MAdul Righ lowe back Sel No Yes
13/12/21 Fishe FAdul Le uppe a m Sel No No
18/03/22 F icka F Adolescen Righ leg Sel Yes Yes 1
Table 1. Summa y o epo ed e en s o insec applica ions based on ideo oo age and abula ed
obse a ional da a as a unc ion o da e, ID (wounded indi idual), sex (M = Male; F = Female), age class o he
wounded indi idual (adolescen s: 8 < x < 16; adul s: x ≥ 1642, wound loca ion, applica o (sel -applied, N = 5;
o applied by a conspeci ic, N = 1), p esence o bys ande pee ing du ing insec applica ion, and a ailabili y o
ideo oo age. The numbe s in he example e en s ma ch he h ee e en s desc ibed in he esul s.
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he insec (wi h only he abdomen isible) on o he wound su ace and picked mul iple ins ances o whi e esidue
om he insec . A one poin , while g ooming he wound wi h he inge s, he insec was pushed aside on o
nea by u , bu she subsequen ly e ie ed i and placed i back on o he wound su ace (Video S1; 01:23 − 01:30).
Du ing he applica ion, whi e esidue was obse ed on se en sepa a e occasions on he lowe lip and humb
(Video S1; imes amps: 00:16, 00:33 [a which ime whi e esidue was also isible on he insec pa ], 00:36, 00:41,
01:04, 01:18), and whi e iscous esidue was also obse ed a ached o he insec pa when disca ded om he
wound (Video S1; 01:43). O hese ins ances, wo demons a ed clea inges ion (Video S1; 00:41, 01:18), whils
in o he cases he esidue was obse ed o adhe e o he lowe lip and chin a ea a e being b ough o he mou h
(Video S1; 00:17 − 00:27, 00:35 − 00:36, 00:38), making i unclea whe he inges ion occu ed. Th oughou he
insec applica ion ac i i y, she p oduced o o acial sounds – ee h clacks and lip smacks43,44.
E en 2 (see Video S2). An adolescen male chimpanzee, Damien, caugh and applied a lying
insec o an open wound on his inne igh cal
The e en in ol ed Damien, an adolescen male, who cap u ed a lying insec wi hin wo a emp s, using a
g asping mo emen on lea es while locomo ing (Video S2; 00:04 − 00:18). Damien p oceeded o p ess he
cap u ed and immobilised insec o his wound on his inne igh cal (app oxima ely < 5cm in diame e ) and
was obse ed e-applying he insec on wo occasions. He held he insec in his mou h o be ween his lips wo
imes be o e eapplying (Video S2; 00:56 − 01:02, 01:06 − 01:08). A he end o he applica ion, he placed he
insec on he edge o his lowe , looked a i be o e disca ding i (Video S2; 01:44 − 01:56). An un ela ed adul
male, Wayne and Damien’s younge adolescen sis e pee ed a he ac i i y a di e en poin s; Wayne s opped
pee ing and walked away be o e he end o he e en (Video S2; 00:22 − 01:58) and E a-James con inued pee ing
un il Damien s a ed o g oom he (Video S2; 01:10 − 02:23).
E en 3 (see Video S3). An adolescen emale chimpanzee, E a James, applied a lying insec
o an open wound on he inne igh cal o he adolescen b o he , Damien
This e en ook place sho ly a e E en 2 (app oxima ely 90s). An adolescen emale and sis e o Damien,
E a James, was obse ed applying an immobilised lying insec o Damien’s open wound (app oxima ely < 5cm
in diame e ). E a James had success ully cap u ed a lying insec by using a quick g asping mo emen on he
nea by lea es (obse a ional eco ding) be o e p oceeding o p ess he insec on Damien’s wound (Video S3;
00:04 − 00:22). Du ing he applica ion, she p essed he insec on Damien’s wound and held be ween he lips
in o he mou h h ee imes, once be o e eapplica ion (Video S3; 00:10 − 00:13) and wice be o e a emp ed
eapplica ion (Video S3; 00:25 − 00:30, 00:51 − 00:54). A e , he second a emp ed applica ion, she walked away
wi h he insec s ill in hand (Video S3; 00:55 − 01:14) es ed nea by and inspec ed he insec be o e disca ding
i (K.K. pe sonal obse a ion). Thei olde adul ma e nal b o he , Rollins, was es ing nea by, who began
Fig. 1. An adolescen emale, F icka, applied a lying insec o a wound on he inne igh high. The wound is
indica ed by he whi e squa e, and he insec , isible as a blue-colou ed objec be ween he humbs, is shown in
he zoomed-in iew wi hin he whi e-bo de ed squa e in he op- igh co ne .
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pee ing a he ac i i y du ing he i s a emp ed eapplica ion un il Damien eposi ioned himsel (Video S3;
00:34 − 00:56).
Discussion
The p esen s udy aimed o p o ide insigh in o insec applica ion o wounds in a di e en chimpanzee subspecies.
We obse ed i e e en s and desc ibe h ee in de ail in ol ing he applica ion o a lying insec on o an open
wound, esul ing in wo sel -di ec ed and one o he -di ec ed applica ion by Eas e n chimpanzees om he Ngogo
popula ion. The obse a ions sugges a gene al sequen ial pa e n in he applica ion o insec s o wounds: i s ,
he insec was cap u ed om nea by ege a ion, such as lea es; second, he immobilised insec o insec pa s
we e applied o he exposed wound su ace using he inge s, o en epea edly, and in some ins ances empo a ily
held in he mou h o be ween he lips be o e being eapplied; and hi d, he insec was ei he disca ded and
emo ed om he wound a ea, wi h no cases o inges ion eco ded. This sequence mi o s ha epo ed in
Cen al chimpanzees, sugges ing a po en ial consis ency in echniques ac oss subspecies. Al hough based on a
small sample size, he indings sugges ha he beha iou is mo e widesp ead han p e iously assumed37, adding
o cu en unde s anding o beha iou al social ansmission and p osocial beha iou in chimpanzees. Hence,
as a ca alys o guide u u e esea ch in o insec applica ions, hei possible eme gence and unc ion, as well as
hei ele ance o he e olu ion o human medical beha iou s, we pos ula e h ee di e en hypo heses ocusing
speci ically on (i) selec i i y o he applied insec , (ii) social ansmission o insec applica ion, and (iii) o he -
di ec ed applica ion as a p osocial beha iou .
Insec selec i i y
All obse ed e en s in ol ed lying insec s a he han o he ypes o insec s such as g ound-dwelling o c awling
insec s, consis en wi h epo s om Cen al chimpanzees37. This epea ed obse a ion may sugges some deg ee
o selec i i y in he ypes o insec s chosen o applica ion. A key ques ion is whe he chimpanzees selec o
hese lying insec s and a speci ic axa o applica ion on hei open wounds and whe he hese applied insec s
ha e medicinal alue.
In humans, lying insec s used o wound healing such as common g een bo le lies (Lucilia se ica a)45–50,
blow lies (Sa conesiopsis magellanica)51,52, as well as honeybees (Apis melli e a)53, and silk mo hs (Bombyx
mo i)54,55, se e se e al opical unc ions, including disin ec ion, an ibac e ial p ope ies, enhanced healing,
issue econs uc ion, and in lamma ion educ ion3. Gi en hese es ablished he apeu ic p ope ies o a ious
lying insec species in humans, chimpanzees may simila ly selec lying insec s wi h medicinal p ope ies o
wound- ela ed applica ions. Howe e , chimpanzees may use hese insec s h ough mechanisms ha also in ol e
sys emic e ec s. E en 1, whe e a whi e esidue was consumed, could indica e sys emic bene i s alongside opical
applica ion. Whe he he inges ion se es a unc ion, and wha mechanisms (solely opical, o bo h opical and
sys emic) migh be in ol ed in he insec applica ion, equi es u he in es iga ion.
Gi en he di icul y o de e mining species-speci ic selec ion by chimpanzees in eal ime, u u e esea ch
could add ess his h ough a wo- old app oach. Fi s , species iden i ica ion can be achie ed h ough collec ing
insec emains om applica ion e en s and iden i ying hem based on mo phological cha ac e is ics o h ough
molecula echniques such as DNA ba coding56. These iden i ica ions should be con ex ualised wi hin he local
insec di e si y, o enable assessmen o whe he chimpanzees p e e en ially selec speci ic species om a wide
ange o a ailable axa. Second, assessing whe he iden i ied insec s ha e medicinal alue and con ain compounds
ele an o wound healing, such as an ibac e ial enzymes, an i-in lamma o y agen s, o o he pha macologically
ac i e subs ances, may be achie ed h ough molecula p o iling57. Compa ing insec s used ac oss di e en
chimpanzee popula ions may o e u he insigh in o whe he insec applica ion e lec s an adap i e esponse.
I insec applica ion aids wound disin ec ion, i may p o ide bo h he apeu ic and p ophylac ic bene i s, wi h
immedia e e ec s (e.g. an imic obial ac ion) and longe - e m bene i s (e.g. quicke eco e y o educed in ec ion
isk), ep esen ing a beha iou wi h clea i ness ad an ages. Such in o ma ion may also p o ide insigh in o he
po en ial social ansmission o he beha iou – i he same o simila insec species a e consis en ly applied –
and in o he possibili y o o he -di ec ed applica ion as a o m o p osocial beha iou , pa icula ly i he applied
insec s ha e medicinal alue (discussed u he in he subsequen sec ions).
Social ansmission o insec applica ion
When del ing in o he insec applica ion i sel and i s gene al sequen ial pa e n, insec applica ions may
necessi a e he ecogni ion and handling o speci ic insec species, which could equi e social lea ning o be
eliably exp essed. One beha iou ha may acili a e he ansmission o insec applica ion and he sequence is
pee ing58,59, as i can p o ide an oppo uni y o acqui e he knowledge o he beha iou . This is suppo ed by
oppo uni ies o obse e conspeci ics, pa icula ly socially ole an indi iduals60, who allow close p oximi y and
epea ed exposu e o hei beha iou 61. Such oppo uni ies a e mo e likely among kin, such as mo he -in an
pai s, bonded indi iduals62, o ole models such as olde indi iduals60. In chimpanzees, ission– usion dynamics
lead o changes in g oup composi ion and pa y size63, esul ing in a ia ion in he a ailabili y o ole models
and a high deg ee o social ole ance60. The acquisi ion o mo e complex beha iou s, such as insec applica ion,
may he e o e depend on ex ended exposu e o expe ienced chimpanzees in ole an con ex s, allowing o mo e
de ailed and eliable ansmission o in o ma ion64.
In his s udy, hi d-pa y pee ing was obse ed du ing insec applica ion e en s (Table1, N = 3 o i e e en s),
sugges ing ha hese in e ac ions may o e oppo uni ies o he social ansmission o he beha iou and i s
sequence. P e ious s udies ha e linked pee ing o he acquisi ion o ool-use beha iou s, p ima ily in eeding
con ex s65–67. In he case o insec applica ion, pee ing may simila ly suppo he ansmission o he beha iou
o aspec s o i s sequence, pa icula ly h ough in e ac ions wi h socially ole an indi iduals o ole models.
The e o e, we sugges ha u u e esea ch pay pa icula a en ion o he p esence o pee ing du ing insec
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applica ion e en s. S udies may examine whe he pee ing p edic s subsequen occu ences o insec applica ion
(ei he sel -di ec ed o o he -di ec ed) by he obse ing indi idual, including simila sequences and insec
selec ion. Fo ins ance, wi h E en 3 which was a ollow-up o E en 2 wi h E a-James who pee ed du ing he
sel -di ec ed applica ion o Damien, soon applied an insec o he b o he Damien’s wound. Ano he addi ional
conside a ion on he insec selec ion, is i hose indi iduals ha pee also applying insec s wi h medicinal alue
(gi en ha he applied insec s possess such p ope ies) o a e hey eplica ing he sequence wi hou selec ing
e ec i e insec s, which could lead o di e en ou comes. S udies should also conside he iden i y o he model
(e.g. kin, olde indi iduals), he ole ance le el (du a ion spen pee ing du ing applica ion wi hou a eac ion by
he model, e.g., mo ing away o di ec ed agg ession), and whe he he pee ing occu ed h oughou he en i e
applica ion o pa ially, as his may in luence lea ning ou comes68.
Insec o he -di ec ed applica ion as a o m o p osocial beha iou
In humans, empa hic esponses o ano he ’s pain a e sugges ed o unde lie he eme gence o p osocial
beha iou 69,70 (see also71. These esponses a e cha ac e ised as au oma ic emo ional eac ions o he a ec i e
s a e o o he s, modula ed by highe -o de cogni i e p ocesses70. A chaeological e idence sugges s ha se e ely
inju ed and disabled Neande hals su i ed o ex ended pe iods, implying ha hey ecei ed ca e and suppo
om conspeci ics72. This kind o o he - ega ding beha iou (i.e., a ge ed helping), po en ially d i en by
empa hic esponses o ano he ’s pain, may ha e oo s in ou las common ances o wi h chimpanzees. Al hough
some schola s a gue ha chimpanzees lack human-like empa hy39,73, o he s udies sugges ha hei beha iou
e lec s empa hy- ela ed endencies, suppo ed by e idence o a ge ed helping in he wild as a o m o epimele ic
beha iou 36,74–80.
The obse ed e en s o o he -di ec ed insec applica ions in Eas e n and Cen al chimpanzees37, con ibu e
o he ongoing deba e on p osocial beha iou s in chimpanzees38,39. Insec applica ion was obse ed be ween
ela ed indi iduals in bo h chimpanzee subspecies: in Eas e n chimpanzees, be ween siblings; and in Cen al
chimpanzees, be ween a mo he and he son. The emaining wo e en s in Cen al chimpanzees in ol ed
un ela ed indi iduals, sugges ing ha , while some ins ances may e lec kin medica ion35, o he s poin owa d a
b oade pa e n o social medica ion35 ha ex ends beyond kinship ies, i shown o ha e a medicinal unc ion.
Howe e , in his s udy, e en s we e eco ded ad libi um; hus, ins ances o o he -di ec ed insec applica ion may
be unde ep esen ed.
Fu u e esea ch could he e o e explo e he possibili y ha o he -di ec ed applica ion is a o m o p osocial
beha iou . Speci ically, i he insec s iden i ied in o he -di ec ed applica ions possess medicinal p ope ies, such
as an imic obial o an i-in lamma o y e ec s, his beha iou may be adap i e and could he e o e quali y as
a ge ed helping. In his con ex , he applica ion o insec s o o he s may p o ide e idence o empa hy- ela ed
capaci ies in chimpanzees, linked o p osociali y. Consequen ly, insec applica ion and he ma ked asymme y
be ween sel - and o he -di ec ed ins ances o e s an oppo uni y o explo e he cogni i e and emo ional
mechanisms ha unde lie ea ly o ms o empa hy and p osocial beha iou .
Insec applica ion as a medica i e beha iou ?
Insec applica ion has no ye been classi ied as medica i e acco ding o he sel -medica ion c i e ia27,81.
These c i e ia ypically include: (1) ecogni ion o symp oms o inju y, (2) use o a subs ance no pa o he
egula die , (3) obse able heal h bene i s ollowing use, and (4) e idence ha he subs ance has biological
o pha macological ac i i y27,81. These c i e ia we e o iginally de eloped o plan use and sel -medica ion.
The e o e, we adap ed hem o help de e mine whe he sel -di ec ed and o he -di ec ed insec applica ion is a
medica i e beha iou :
(1) Recogni ion o wounds.
(2) The iden i ied insec and i s de i ed ma e ial p o ide li le o no nu i ional bene i i consumed.
(3) Obse able wound healing ollowing use.
(4) E idence ha he applied insec has biological o pha macological ac i i y.
C i e ia (1), (2), and (4) can be in es iga ed ela i ely easily h ough beha iou al obse a ion, die a y analysis,
and chemical o mic obiological es ing. In con as , c i e ion (3) may pose mo e challenges, as healing ou comes
can be a ec ed by a ious con ounding ac o s, including he se e i y and loca ion o he wound, indi idual
a ia ion in immune esponse, and en i onmen al condi ions. Ne e heless, i c i e ia (1), (2), and (4) a e clea ly
sa is ied, his may p o ide s ong e idence in a ou o classi ying insec applica ion as a medica i e beha iou .
Conclusion
This s udy p esen s he i s documen ed e idence o insec applica ion in Eas e n chimpanzees, ex ending a
beha iou p e iously epo ed only in Cen al chimpanzees. This inding expands he known epe oi e o wound
ca e in chimpanzees and sugges s ha insec applica ion may be mo e p e alen han p e iously ecognised. The
documen a ion o bo h sel -di ec ed and o he -di ec ed insec applica ions poin s o i s po en ial ole wi hin
he b oade medicinal beha iou al epe oi e o chimpanzees, o e s insigh in o hei cogni i e complexi y,
and in oduces a c ucial candida e beha iou o unde s anding he e olu ion o human medical beha iou s.
Mo e b oadly, hese indings highligh he impo ance o p ese ing wild chimpanzee popula ions, no only
o hei con inued su i al bu also o main aining he beha iou al di e si y o which insec applica ion may
meaning ully con ibu e82.
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Me hods
S udy si e and subjec s
The indi iduals obse ed we e membe s o he Ngogo chimpanzee popula ion li ing in he Kibale Na ional
Pa k in Uganda. A he ime o his s udy, was di ided in o wo dis inc communi ies: Cen al and Wes e n83,
and he membe size o he popula ion a ied be ween 180 and 220 indi iduals. Al hough o iginally a single
communi y wi h blu ed socio-spa ial bounda ies be ween h ee subg oups (Wes e n, Cen al, and Eas e n), a
se ies o agonis ic e en s led o a pe manen spli in 2017, esul ing in he o ma ion o wo dis inc communi ies
wi hin he Ngogo popula ion83. The s udy a ea is app oxima ely 30 km2 and consis s o mos ly ma u e and
egene a ing o es ansi ional be ween lowland and mon ane mois e e g een o es s, bu also smalle a eas o
swamp o es 84.
Beha iou al obse a ions
F om No embe 2021 o Ma ch 2022, all e en s in ol ing insec applica ion we e collec ed by K.K. and D.A.F.
ad libi um85. All occu ences o open wounds we e eco ded, and i possible, he cause o he wound was
documen ed. Open wounds included punc u es, lesions, lace a ions, and small cu s. Healed sca s and scabs we e
excluded. Sel -di ec ed and o he -di ec ed insec applica ions we e eco ded digi ally using he Cybe T acke
p og amme ( e sion 3.514) on a wa e p oo sma phone. When possible, ideo oo age o he e en was eco ded
using a digi al high-de ini ion came a (Sony FDR-AX53 4K). The ID o he wounded indi idual and applica o ,
beha iou al sequence o insec applica ion, wound loca ion, and in ol emen o pee ing (i.e., close- ange and
sus ained obse a ion o a conspeci ic’s ac i i ies86 we e no ed o each obse ed e en .
E hics s a emen
The esea ch conduc ed on he chimpanzees was s ic ly non-in asi e and obse a ional, and was classi ied
as a non-animal expe imen unde Sec ion V, A icle 7 o he Ge man Animal Wel a e Ac (25 May 1998). I
was app o ed by he Ugandan Wildli e Au ho i y ( e : COD/96/05), he Ugandan Na ional Council o Science
and Technology (no. NS295ES), he Make e e Uni e si y Biological Field S a ion, and he Ngogo Chimpanzee
P ojec . The esea ch adhe ed o he ASAB Guidelines87 and he IUCN Bes P ac ice Guidelines88, bo h o which
se in e na ionally ecognised s anda ds o e hical, non-in asi e beha iou al ield esea ch on animals, including
p ima es. These s anda ds guided all aspec s o ieldwo k, including p e- ieldwo k qua an ine p ocedu es, he
use o su gical masks, and main aining a dis ance o app oxima ely 10m om he chimpanzees.
Da a a ailabili y
Da a is p o ided wi hin he manusc ip and ideo oo age o epo ed e en s is a ailable a he ollowing epos-
i o y: h ps: //o s .io/a j5 8 /? i ew_only=575674748 c254725b 91a9a b2542cd .
Recei ed: 19 June 2025; Accep ed: 18 Augus 2025
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Acknowledgemen s
This wo k was unded by he Deu sche Fo schungsgemeinscha (DFG) Ge man Resea ch Founda ion, p ojec
numbe 274877981 (GRK-2185/1: DFG Resea ch T aining G oup Si ua ed Cogni ion), which suppo ed K.K.,
and he Eu opean Resea ch Council (ERC), EU-Consolida o g an (772000, Tu nTaking) unde he Eu opean
Union’s Ho izon 2020 Resea ch and Inno a ion P og amme awa ded o S.P. We a e g a e ul o he di ec o s John
C. Mi ani, Ke in Lange g abe , and Da id Wa s o hei app o al o s udy he chimpanzees o he Ngogo Chim-
panzee P ojec . We a e hank ul o he Ugandan Wildli e Au ho i y, he Make e e Uni e si y Biological Field
S a ion, and he Ugandan Na ional Council o Science and Technology o hei pe mission o conduc esea ch
a he Kibale Na ional Pa k. We hank Samuel Angedakin o assis ance wi h he necessa y adminis a ion and
pe mi s. We a e indeb ed o Sebas ian Rami ez-Amaya o his suppo and e o in assis ing wi h da a collec-
ion. Las ly, we hank he membe s o he Compa a i e BioCogni ion Resea ch G oup o ui ul discussions
and adminis a i e suppo , and he chimpanzees a Ngogo, whose ole ance made his esea ch possible.
Au ho con ibu ions
Concep ualisa ion: K.K., and S.P.; Manusc ip shaping: S.P.;Da a Cu a ion: K.K.; Fo mal Analysis: K.K.; Funding
acquisi ion: S.P.; In es iga ion: K.K., and D.A.F.; Resou ces: S.P.; W i ing – o iginal d a : K.K.; W i ing – e iew-
ing and edi ing: K.K., D.A.F., A.M., and S.P.; Visualisa ion: K.K.; Supe ision: S.P.
Funding
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