Pan Africa NewsThe Newsletter of the Committee for the Careand Conservation of Chimpanzees, and the Mahale
Wildlife Conservation Society
ISSN 1884-751X (print), 1884-7528 (online) mahale.main.jp/PAN/

DECEMBER 2017
P. A. N. EDITORIAL STAFF
Chief Editor:
Kazuhiko Hosaka, Kamakura Womenʼs University, Japan
Deputy Chief Editor:
Michio Nakamura, Kyoto University, Japan
Associate Editors:
Christophe Boesch, Max-Planck Institute, Germany
Jane Goodall, Jane Goodall Institute, USA
Tetsuro Matsuzawa, Kyoto University, Japan
William C. McGrew, University of St. Andrews, UKJohn C. Mitani, University of Michigan, USA
Vernon Reynolds, Budongo Forest Project, UK
Yukimaru Sugiyama, Kyoto University, Japan
Richard W. Wrangham, Harvard University, USA
Takeshi Furuichi, Kyoto University, Japan
Editorial Secretaries:
Noriko Itoh, Kyoto University, Japan
Koichiro Zamma, Nagano College of Nursing, Japan
Eiji Inoue, Toho University, Japan
Takuya Matsumoto, Research Institute for Humanity and Nature,Japan
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VOL. 24, NO. 2

Contents<NOTE>A Female Bonobo Sleeping on the Ground afterDaytime Birth and its Implications
Heungjin Ryu

9<NOTE>
Lonoa: The Establishment of a Permanent Field
Site for Behavioural Research on Bonobos in the
Kokolopori Bonobo Reserve
Martin Surbeck, Sally Coxe, &Albert Lotana Lokasola

13<NOTE>
Termite Fishing by Mahale Chimpanzees: Revisited,
Decades Later
Alejandra Pascual-Garrido
15<NOTE>An Old Female Bonobo Carried a Dead Red-Tailed
Monkey for over a Month
Kazuya Toda, Nahoko Tokuyama, &Takeshi Furuichi
19<BOOK REVIEW>
Cannibalism: A Perfectly Natural History (By Bill
Schutt)William C. McGrew
21<NOTE>A Female Bonobo Sleeping on the
Ground after Daytime Birth and
its Implications
Heungjin Ryu1,2,3
1 Primate Research Institute, Kyoto University, Japan
2 Research Institute of Eco-science, Ewha Womans University,
Republic of Korea
3 National Institute of Ecology, Republic of Korea(Email: ryu.heungjin.26v@st.kyoto-u.ac.jp)

INTRODUCTION

Reduction of predation risk is thought to be one of
the main benefits that animals derive from living in agroup (Alexander 1974) and a major driving force in theevolution of group-living in primates (Hill & Lee 1998).

9Pan Africa News, 24(2), December 201710

Figure 1. Ot with her newborn infant (OS) on the ground. Ok and Fa looking at OS. Fklying about 2 m from the other three and looking back at human observers.

For bonobos and chimpanzees, the combined effects oflarge body size and group living may mean that predation
risk is relatively low compared with many other primate
species, but some reports have suggested that lions andleopards can have a heavy impact on them (Boesch 1991;
Tsukahara 1993; D’Amour et al. 2006). Among chimpanzees there is an additional risk from conspecific killing,which is probably responsible for deaths of more infants
and adults than predation itself (Williams et al. 2008).
Although they are closely related, bonobos are quite
different from chimpanzees in terms of conspecific aggression and female–female relationships (Furuichi 2011).
Neither direct evidence of infanticide nor conspecifickilling has been observed in wild bonobos. Female chimpanzees are less tolerant and compete with each otherfor local resources, probably as a means to increase theirreproductive success (Pusey & Schroepfer-Walker 2013).
Female bonobos, on the other hand, often engage in affiliative interactions such as grooming and homosexualinteractions called genito-genital rubbing (Kuroda 1980;
Ryu et al. 2015) and may support each other in response
to aggression from males (Tokuyama & Furuichi 2016).
Female bonobos keep this high sociality even during one’s
parturition (Douglas 2014). This friendly social environment appears to influence many behavioral characteristics
of female bonobos.
Here I report a case of a prime parous female bonobo
giving birth in the daytime at Wamba in the Luo Scientific
Reserve, Democratic Republic of the Congo. The subject
female, called Otomi (Ot) gave birth on a daybed on April
5th, 2015. After giving birth, Ot came down to the groundand fell asleep while her nearest neighbor Fuku (Fk) was
sitting about 2 m from her. I will discuss how tolerant andsupportive group characteristics can be related with behavioral characteristics of female bonobos compared withthat of female chimpanzees.

METHODS AND BACKGROUND

E1 group was originally one of the two subunits ofE group which had been habituated by 1976 (Furuichi etal. 2012). Since then, E1 group has been studied for over
40 years at Wamba, with a break of 6 years from 1996
to 2002 because of the civil war in DR Congo. When researchers resumed the field study in 2002, all membersof E1 had been re-identified and no artificial provisioning
has been conducted since that time (Furuichi et al. 2012).
I followed E1 group from dawn to dusk (bed to bed)
with two field assistants. We recorded all occurrences ofrare behaviors, such as aggression and copulation, as wellas membership of the party every hour using the 1-hourparty method (Hashimoto et al. 2001).
Otomi (Ot) was one of two females that immigrated
into E1 from an adjacent group in 2008, at which time she
was estimated to be 11 years old (Sakamaki et al. 2015).
Ot successfully gave birth to her first offspring in January
2011, and it was named Otoko (Ok; ♀). Ok had grown up
and become an independent individual by 2015. Ot hada still-birth in March 2014 when Ok was 3 years and 2
months old. Ot became pregnant again 5 months after thestill-birth. This time Ot successfully gave birth on April 5,
2015, and this offspring was named Osamu (OS; ♂).
Two months prior to Ot’s immigration, another immigrant female Fuku (Fk) joined E1 group. Fk also gave
a birth in January 2011, and her offspring was named Fua(Fa; ♀). The relationship between Ot and Fk was stable
and strong in 2015 when the observations in this reportwere made.

OBSERVATIONS

At 05:31–06:00 h, April 5th, 2015, Ot was found without newborn infant at the bed site. E1 group moved into
the secondary forest, where Musanga trees (Musanga cecropioides) are most abundant.
At 07:45 h, Ot stayed in a daybed and I could not detect any particular movement nor vocalization (focal sam-Pan Africa News, 24(2), December 2017
pling). The only other daybed in the same tree was Ok’s,(about 2 m distance). Two more daybeds were around at
7 m from Ot’s bed (individuals in them were not identified). I left the place at 08:05 h when the focal sampling
was over.
At 08:45–08:50 h, Ot urinated on the daybed, and left.
The urine found on the leaves of shrubs below included
some drops of blood. Ot was carrying a newborn infant(OS) with umbilical cord (around 5 cm) still attached. Ot
moved into a bushy place and lay on the ground. Ok followed Ot.
At 08:51–09:22 h, Fk and Fa appeared at the place
where Ot and Ok were. Fk stopped about 2 m from Ot.
Fa and Ok approached within arm’s reach of Ot to see OS(Figure 1, Video 1 available online at http://mahale.main.
jp/PAN/24_2/24(2)_01.html). Ot sometimes shooed themaway when they got too close to OS (Figure 2). Fk waswatching Ot from about 2 m away and did not get closer.
While taking a rest on the ground, Ot fell asleep and tried
to open her eyes (Figure 3 and Video 1). I left with the
trackers to avoid disturbing them anymore.
Ot and her two offspring were then absent from thegroup until April 13, 2015 (8 days). TN (an adult male;over 40 years old) was also missing from 7th to 13th andit may be that TN ranged with Ot and her two offspring.

DISCUSSION

After giving birth, a bonobo mother might be expected to experience sleepiness, as shown in the current
report. However, sleeping on the ground will increase therisk of being attacked by predators or conspecific individuals. Therefore, it is hard to imagine that a female of a solitary primate species could fall asleep on the ground witha newborn infant. This might be also true for a female
living in an intolerant social environment, such as that ofchimpanzees. Female chimpanzees increase vigilance andself-directed behaviors when there are more members in

11
close proximity (Kutsukake 2003, 2006). Moreover, otherconspecifics can seriously harm one’s newborn infant inchimpanzees (Goodall 1977; Nishie & Nakamura 2018).
However, if the risk of predation or harmful contact byconspecifics is very low, females might not need to maintain a high level of vigilance while they are with othergroup members as described in the current report.
Female bonobos maintain close social relationships
with each other and support one another in agonistic interactions. These close social relationships and agonistic
support enable females to maintain high social status as
they are able to counter any attempts by males to dominate them (Parish 1996; Tokuyama & Furuichi 2016). The
current report is in line with these findings. If a female
bonobo with other group members can benefit from theirprotection from predation and conspecific harassment, theneed for maintaining vigilance will be low. Douglas (2014)
also suggested that females get benefit of protection andrelief from others during the parturition. Therefore, thecohesive grouping pattern and tolerant female–female
relationship might be the medium that makes female
bonobos relaxed, even at times when they are relatively
vulnerable, such as immediately postpartum.
The presence of human observers is unlikely to havebeen a cause of Ot’s post-partum sleeping. Even thoughbonobos of E1 group are well habituated, when humanobservers accidently get too close to bonobos (within
about 5 m), they usually move away from the human observers. Females are particularly sensitive to accidental
approaches by human observers. Therefore, it seems most
likely that the presence of Fk and the stable relationshipbetween them facilitated post-partum sleeping by Ot. It
is unlikely that Fk had tracked Ot’s movements until she
gave birth. However, given that frequent proximity andaffiliative interactions between Ot and Fk, Fk was most
likely to be the individual whom Ot would encounter for
the first time after giving birth.

Figure 2. Ot pushing Ok away from the newborn infant, OS.

Pan Africa News, 24(2), December 2017

12

Figure 3. Ot has fallen asleep on the ground with her newborn infant (OS), while Fk, Ok, Fa
stay around her within 2 m.

Nishie & Nakamura (2018) suggested that the temporal leaving of females after giving birth functions as“maternity leave” in chimpanzees to avoid the risk of
infanticide. In the current study group (E1), females withnewborn infants also range for one to two weeks alone, oronly with their other offspring, immediately after giving
birth. However, after one or two weeks, females usually
join a party and range with group members. This is probably because the mother with a newborn infant can findfruiting trees and avoid predation risk more efficiently
by ranging with other group members. Together with thefact that no infanticide has ever been reported in bonobos,
the temporal leaving of female bonobos after parturition
is more likely related with the cost of ranging with othergroup members rather than the risk of infanticide.
The tolerant group characteristics of bonobos mightalso allow female bonobos to exploit the benefit of groupliving more than female chimpanzees can do. This highlights the impact that social environment can have on
behavioral characteristics of females. Further comparative studies on difference of behavioral characteristics ofthe two Pan species will not only help us to understandthe Pan species, but may also improve our understandingof how social characteristics of various human societies
influence well-being and behavioral characteristics of theindividuals in them.

ACKNOWLEDGEMENTS

I thank research assistants at Wamba and the ResearchCenter of Ecology and Forestry for their great contribution at
the study site. I also thank Dr. Takeshi Furuichi, Dr. Tetsuya
Sakamaki for their support. I also thank Dr. Hitonaru Nishie
and Dr. Kazuhiko Hosaka for reviews on the manuscript.
Finally, I thank Dr. David A. Hill for comments on the manuscript and English editing. This study was permitted by the
Ministry of Scientific and Technological Research of the D.R.
Congo (MIN.ESURS/SG-RST/026/2014) and funded by theJapan Society for the Promotion of Science (JSPS) Asia-Africa
Science Platform Program (2012–2014 to Furuichi).

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Douglas PH 2014. Female sociality during the daytime birthof a wild bonobo at Luikotale, Democratic Republic of theCongo. Primates 55:533–542.
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Furuichi T 2011. Female contributions to the peaceful natureof bonobo society. Evol Anthropol 20:131–142.
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Furuichi T, Idani G, Ihobe H et al. 2012. Long-term studies
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Hashimoto C, Furuichi T, Tashiro Y 2001. What factors affect the size of chimpanzee parties in the Kalinzu Forest,
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Parish AR 1996. Female relationships in bonobos (Pan paniscus): Evidence for bonding, cooperation, and female dominance in a male-philopatric species. Hum Nat 7:61–96.
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Sakamaki T, Behncke I, Laporte M et al. 2015. Intergrouptransfer of females and social relationships between immigrants and residents in bonobo (Pan paniscus) societies. In: Dispersing Primate Females: Life History andSocial Strategies in Male-Philopatric Species. Furuich T,
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Lonoa: The Establishmentof a Permanent Field Site for
Behavioural Research on Bonobos
in the Kokolopori Bonobo
Reserve13
have been traditionally protected by a taboo of the local Bongando people against hunting them (Lingomo& Kimura 2009). The management of the reserve isconducted by the local population and the conservation
initiative “Vie Sauvage” (VS), supported by the Bonobo
Conservation Initiative (BCI).
Efforts to follow bonobos in the northern part of thereserve, near the village of Yetee, were initiated in 2005
by VS and BCI, which had been committed to establishing longitudinal research on bonobos and had commenced
ongoing support of local people to track the Ekalakala
bonobo community (Figure 2). In 2010, the tracking
extended to a second, neighboring bonobo community,
Nkokoalongo. Until 2016, both communities had been
followed on a more or less regular schedule, but were
subject to only a limited amount of formal scientific work.
Dr. Alex Georgiev did some work in the ranging area of
the Ekalakala community, collecting fecal samples during 2006 and 2007. These samples were included in a
number of publications on malaria and micro biome (Liuet al. 2010; Liu et al. 2014; Ahuka-Mundeke et al. 2016;
Moeller et al. 2016; Wroblewski et al. 2017). Dr. Deborah
Moore commenced behavioural ecological research at thesite in 2014 with the intention of establishing a longitudinal study, but unfortunately was curtailed after 4 months.
During her stay, she reorganized the habituation procedure for both communities and made some initial behavioural observations, including an incidence of maternalcannibalism (Tokuyama et al. 2017).
After MS’s reconnaissance trip in 2015, he established a permanent research camp named after the nearby
river, Lonoa (N0.41716°, E22.97552°), in the beginningof 2016 with the help of the Max Plank Institute, VS andBCI. The camp is situated approximately 4 km from thenearest village, Yetee (Figure 1). A network of existing
paths and standardized transects allows access to an areaof 50 km², which comprises large parts of the ranges of
the two habituated bonobo communities. The study area
consists mainly of primary heterogeneous forest with very
few swampy areas. The bonobos of the Ekalakala community also range occasionally in secondary forests close
to the local settlements. An initial botanical survey todescribe the forest types and composition has been con-Martin Surbeck1,2, Sally Coxe2, &Albert Lotana Lokasola2,3
1 Max-Planck-Institute for Evolutionary Anthropology,
D-04103 Leipzig, Germany
2 Bonobo Conservation Initiative, 2701 Connecticut Ave. NW,
Suite 702, Washington DC 20008, US
3 Vie Sauvage, Avenue Pangi 5, commune Ma Campagne,
Kinshasa, DRC(E-mail: surbeck@eva.mpg.de)

The Kokolopori Bonobo Reserve was officially established by the Democratic Republic of the Congo (DRC)
government in May 2009 as a community-based nature
reserve encompassing over 4,850 km² (Almquist et al.
2010; Figures 1 and 2). Locally initiated and managed,
it harbors a large population of bonobos, who are andFigure 1. Active bonobo research sites with at least semihabituated bonobo communities.

14

Pan Africa News, 24(2), December 2017
ture individuals. The Nkokoalongo community consistedof 10 adult and subadult males, 15 parous and 1 nulliparous female, and 15 immature individuals.
While research on bonobos at Kokolopori is still in an
early phase, the site has great potential not only to explore
behavioural variations within the species due to a different habitat as compared to the long-term research sites ofWamba (Furuichi et al. 2012) and LuiKotale (Hohmann& Fruth 2003), but also to address questions concerning
relationships between communities with largely overlapping areas. Given the well-established positive impactof long-term research sites on conservation efforts (e.g.,
Pusey et al. 2007, Campbell et al. 2011), the presence of along-term research project in the protected site gives added hope that the bonobo population is relatively safe frompoaching for bush-meat and timber exploitation. However,
due to the proximity to human settlements, there is a needfor constant vigilance.

Figure 2. Kokolopori Bonobo Reserve. Approximate
home ranges of the two study communities areshown in dotted circles.

ducted in collaboration with the University of Kisangani,and a floristic comparison with other bonobo study sites ispart of an ongoing study.
While hunting is not allowed in large parts of thestudy area, the use of the forest is shared with the local people, whose primary activities there are collecting
caterpillars and mushrooms in their respective seasons.
Due to the close proximity of human settlements, animal densities are rather low compared, for example, tothe LuiKotale bonobo study site at the border of Salonga
National Park (Hohmann & Fruth 2003). However, initial
results of camera-trapping indicate the occurrence of several species of duikers including blue duiker (Cephalophus
monticula), black-fronted duiker (Cephalophus nigrifrons), bay duiker (Cephalophus dorsalis) and Peter’s
duiker (Cephalophus callipygus), and monkeys including
red-tailed monkeys (Cercopithecus ascanius), Wolf’smonkey (Cercopithecus wolfi) and De Brazza’s monkey(Cercopithecus neglectus). Furthermore, there are raresightings via camera traps of red riverhog (Potamochoerus
porcus), sit at u nga (Traelaphus spekei), a ardvark(Orycteropus afer), tree pangolin (Phataginus tricuspis),and golden cat (Felis aurata). Combining the results fromthe ongoing camera trap survey with the sightings madeduring an US Fish and Wildlife Service funded survey
conducted in 2011–2012 will facilitate better estimates ofwildlife diversity, abundance and its changes.
After fieldwork began in 2016, a team of at least two
people has followed each of the two neighboring communities daily from nest to nest. From March 2016 to July
2017, the average number of monthly nest-to-nest follows
was 28 for Ekalakala, and 27 for Nkokoalongo. At thisstage, the individuals of both bonobo communities arewell habituated to the presence of observers, allowing for
behavioural studies that require close range observationsof focal individuals. All individuals in both communities were identified and named by the end of 2016. The
Ekalakala community consisted of 3 adult and subadultmales, 5 parous and 1 nulliparous females, and 5 imma-

ACKNOWLEDGEMENTS

We thank the Max Planck Society for providing the funding for the research at Lonoa; Bienvenu Mupenda, Claude
Baombo, Leonard Nkanga, Axel Martìnez Ruiz, Maëlle
Lemaire, Rodolphe Violleau and Claudia Wilke for their help
in local negotiations, constructions and habituation of thebonobos; all members of BCI and VS for their helpful support
during various stages of the project. Nearly 15 years of tracking, monitoring, and habituating the bonobos by the local teams
supported by BCI and VS enabled gave us an enormous headstart. Our special thanks to the late Deborah Moore for her efforts to initiate a long term research project on the bonobos at
Lonoa; we wish she were still there to work with us.

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Termite Fishing by Mahale
Chimpanzees: Revisited, Decades
Later
Alejandra Pascual-Garrido
School of Archaeology, University of Oxford, UK(E-mail: alejandra.pascual-garrido@rlaha.ox.ac.uk)

INTRODUCTION

Jane Goodall’s discovery in 1960 that wild chimpanzees at Gombe Stream National Park in Tanzania
regularly make and use tools to fish for termites was ofgreat importance, as scientists thought at the time thathumans were the only species to make and use tools(Goodall 1964). On hearing of Goodall’s observation, her
mentor, the paleoanthropologist Louis Leakey, famously
remarked: “Now we must redefine ‘tool’, redefine ‘man’,or accept chimpanzees as ‘humans’”. But did only Gombe
chimpanzees do this?
Since then, evidence of termite fishing has been de-scribed in many wild populations of chimpanzees, and is
now recognized as one of the most widespread forms ofchimpanzee technology (McGrew 1992). The first evidence of termite fishing by the chimpanzees of Bilenge(B group) that inhabit Mahale Mountains National Park in
Tanzania dates to 1975. At that time, a local field assistant to the Japanese team of primate researchers by chance
discovered recently abandoned plant tools close to a termite mound (Nishida & Uehara 1980). In the following
years, more ad libitum records followed. These findings
suggested that chimpanzees of B group habitually fish fortermites, using 16 plant species to manufacture implements made (mostly) from bark, but also from sedges,
vines, twigs and leaves (Nishida & Uehara 1980; McGrew& Collins 1985).
Over the last three decades, termite fishing of B grouphas remained unstudied, as research focus shifted to thehabituated neighboring Mimikire (M) group (Nakamura
et al. 2013).
However, my recent research provides evidence thatthe Bilenge chimpanzees have continued with their termite fishing tradition, providing yet another example ofthe long-term maintenance of material culture traditions
in our closest living relatives.

METHODSThe unhabituated chimpanzees (Pan troglodytes
schweinfurthii) of B group live around Bilenge (S 6°2′,
E 29°44′; 772–1550 m altitude) in the Mahale Mountains
National Park, western Tanzania (Nishida 1968; McGrew& Collins 1985). Bilenge is open grassy woodland, mostly
miombo, with narrow strips of vine tangle and valley
forests with broad hilltops rising from the coastal plain(Collins & McGrew 1988).
Earlier studies indicated that termite fishing at
Bilenge is seasonal, with the most productive months in
November and December coinciding with the annual reproductive and dispersal cycles of the termites (McGrew& Collins 1985). My fieldwork, aided by Tanzanian fieldassistants, thus encompassed three such periods (25Nov–14 Dec 2014; 20 Oct–8 Nov 2015; 21 Nov–8 Dec
2016). During reconnaissance surveys, we first searched
for mounds that had been targeted previously by chim-

Table 1. Plant species used by chimpanzees of B group at Mahale Mountains
National Park to source termite fishing tools.

Species
Uvaria angolensis
Uvaria sp. A of FTEA
Monanthotaxis buchananii
Maerua holstii
Garcinia buchananii
Allophylus congolanus
Paullinia pinnata
Mimusops kummel
Grewia stolzii
Saba comorensis
Unidentified
Total

Tools (n) Individual plants (n) Parts sourced* (n)
−
8
2
2
1
−
−
−
4
−
8
25

24
9
3
1
1
3
1
1
3
1
−
47

* Individual plants sometimes provided more than one sourced part.

33
10
6
1
1
3
1
1
4
1
−
61

Pan Africa News, 24(2), December 2017

16

a

b

Figure 1. Plant (Monanthotaxis buchananii) recently sourced by Mahale chimpanzees for bark. (a–b) Detached barkleft on source plant as chimpanzees removed material to manufacture tools. Photo: Alejandra Pascual-Garrido.

panzees, defined by one or more of the following criteria(cf., Pascual-Garrido et al. 2012; Almeida-Warren et al.
2017): (a) tools, fragments or modified raw materials were
present; (b) nearby plants had been sourced for raw material, indicated by, e.g., broken stems or branches or parts
removed; (c) abandoned tools could be retrospectively fitted to their sources.
Measurements included tool characteristics such aslength, diameter, type of raw material, approximate age(estimated number of days since tool was used), as well
as source species of vegetation, location of sourced parts
within the plant (i.e., branch, stem) and height and diameter of stem or branch at point of material removal. Termite
specimens from targeted mounds fixed in ethanol 80%
were later identified by Rudolf Scheffrahn, University of
Florida, USA. Samples from source plants were processedat camp with a plant drier for later identification by Frank
Mbago, Botany Department, University of Dar es Salaam,
Tanzania.

RESULTS

B group members have maintained their termite fishing tradition for more than four decades, as indicated by
2014–2016 records of 25 tools and 47 individual source
plants, some of them sourced multiple times, comprisingat least 10 species (Table 1). Source plants were foundnear 14 mounds (Figure 1 a–b), with 5 of these identified
as Macrotermes michaelseni. One termite mound yielded
tools on four dates (1 Dec 2014; 8 Dec 2014; 26 Nov 2016;a

28 Nov 2016).
Recovered tools (n = 25) were traced back to 5 plant
species, with 47% manufactured from an individual of
Uvaria sp. A of FTEA, followed by Grewia stolzii (23.5%),
with 3 tools manufactured from a single plant. Maerua
holstii and Monanthotaxis buchananii were equally represented in the tool assemblage (11.7%), while Garcinia
buchananii was the least represented (5.8%). All parts detached from plant sources (n = 61) were from bark extractions, with the majority sourced from Uvaria angolensis(54.0%) and Uvaria sp. A of FTEA (16.3%) (Table 1).
All recovered tools were less than 24 h old, and madeexclusively from bark (Figure 2 a–b). The mean length oftools was 38.5 cm (SD = 10.3, n = 25, range: 21.1−65.2),
while mean diameter measured at medial point (approximately half-way along the tool) was 3.1 mm (SD = 1.0,
n = 25, range: 1.5−5.1). Chimpanzees sourced the raw
material (n = 58) mainly from branches (96.5%) and only
rarely from stems of plants (3.4%). The vast majority(91.3%) was removed from up to 1.5 m above ground, at amean height of 92.7 cm (SD = 50.8, n = 58, range: 5−280)(Figure 3). The mean diameter of the stem measured at
the point from where the tool material was removed was
8.5 mm (SD = 2.1, n = 58, range: 15.5−5.3).

DISCUSSION

This recent study shows that termite fishing has been
maintained by the B group of Mahale chimpanzees for atleast 42 years. While the physical characteristics of theb

Figure 2. Termite fishing implements abandoned by chimpanzees after foraging at a mound of Macrotermes
michaelseni (1 Dec 2014). (a) Bark fishing tools in situ, with fresh leaves removed during the manufacturing
scattered on ground, (b) Tool assemblage reflecting a fishing event (1 Dec 2014). Photo: Alejandra Pascual-Garrido.

Pan Africa News, 24(2), December 201717

Table 2. Tools, plant source species and targeted prey associated with termite fishing by chimpanzees of B group
at Mahale Mountains National Park. Bold = plant species sourced for raw material in more than one study.

Averagetool length(cm)
54.6 (range
21.7−125.6)

Averagetool width(mm)

TermiteSpeciesTime ofcollection

Source−M. herus

Oct–Nov
1978

Uehara
1982

133

−

−

M. herus

Nov–Dec
1979

Nishida& Uehara
1980

Bark, sedge,vine, twig, leaf

290

37.7 cm(SD = 14.7)

4.6 mm(SD = 2.0)

M. herus

MainlyNov–Dec
1982

McGrew& Collins
1985

Bark

25

38.5 cm(SD = 10.3)

3.1 mm(SD = 1.0)

M.
michaelseni

Dec2014/
Nov2016

Current
study

Plant Species

Type ofmaterial

Tools
(n)

−

Bark?

97

Bark
Bark
Bark
Bark
Bark
Vine
Vine
Twig
Twig
Grass, bamboo
or sedge
Grass, bamboo
or sedge
Grass, bamboo
or sedge

Uvaria angolensis
Bauhinia petersiana
Grewia platycada
Grewia mollis
Acalypha sp.
Rhynchosia luteola
Tylophora sp.
Rothmania manganjae
Milletia angustidentata
Olyra latifolia
Hyparrhenia rufa
Cyperus diffusus
Acalypha chirindica
Artabotrys monteirode
Bauhinia petersiana
Carpolobia alba
Cyphostemma sp.
Cyperuspseudoleploclaudus
Monanthotaxis
buchananii
Uvaria angolensis
Uvaria sp. A of FTEA
Saba comorensis
Maerua holstii
Garcinia buchananii
Allophylus congolanus
Paullinia pinnata
Mimusops kummel
Grewia stolzii

tools, such as length and width, are similar with thosereported decades ago (McGrew & Collins 1985), somedifferences emerged in relation to the species of termite
prey and the type of material and species used for tool
manufacture.
While bark also was predominantly used in the past,other materials such as twig, sedge, vine, grass, bamboo,
and leaf also were employed (Nishida & Uehara 1980;
McGrew & Collins 1985). That only tools made from bark
were recovered in this study may reflect a smaller sample size compared to previous tool assemblages (Nishida& Uehara 1980; McGrew & Collins 1985) (Table 2).
Similarly, that only sources with evidence of bark extraction were found might be linked to the high fidelity that“scars” associated to these type of material extractions
preserve on the source plant, compared to those associatedwith the detachment of other material such as twig, vineand grass, which hardly ever preserve (Pascual-Garrido,
submitted).

Alternatively, members of B group may have become
exclusive bark users in recent decades. If so, then frequency differences in raw materials may reflect changes
in their availability (McBeath & McGrew 1982). For example, the cessation of annual burning practice during
the last decades might have allowed natural regenerationof woody vegetation near mounds (Collins & McGrew
1988). It also may be that historical variation in tool material preference is responsible (Hobaiter et al. 2014).
Chimpanzees show material selectivity for manufacturing certain type of tools, including termite fishingprobes (McBeath & McGrew 1982; Sanz & Morgan 2007;
Almeida-Warren et al. 2017), a skill possibly transmitted
across generations through social learning, such as via
tool transfers (Sanz et al. 2014; Musgrave et al. 2016).
However, without behavioral evidence of cultural transmission of tool material preference for this study community, this hypothesis remains untested. That measured
length and width of the recovered tools seem to havePan Africa News, 24(2), December 201718

Figure 3. Frequency distribution of heights of sourced plant parts at point of
detachment over 50-cm classes.

been largely maintained for decades may be the productof socially learned skills (Sanz et al. 2014; Musgrave etal. 2016). Alternatively, this could simply be the result of
ecological circumstances such as constancy of raw materials (McBeath & McGrew 1982).
From the total of source species recorded in thisstudy, only Uvaria angolensis has been used in thepast, also as a source of bark (Nishida & Uehara 1980).
However, Artabotrys monteiroae, Paulinnia pinnata andGrewia sp. were used by the formerly neighboring, but
now extinct, K group to fish for Pseudoacanthotermes
spiniger (Uehara 1982). Interestingly, Saba comorensis,
Grewia sp, Monanthotaxis buchananii, Uvaria angolensis, and Uvaria sp. A of FTEA are currently sourced fortermite fishing bark tools by Issa Valley chimpanzees
living 85 km to the east (Almeida-Warren et al. 2017),
while Uvaria angolensis, Allophylus congolanus and Saba
comorensis constitute sources for termite fishing implements at Gombe, 200 km to the north (Pascual-Garrido,
submitted).
Different communities of chimpanzees using thesame species of plants for tool material could result fromtheir relative abundance. Alternatively, these species
might have certain physical characteristics that makethem ideal as tool raw material, i.e., optimal flexibility(Teleki 1974). This, in turn, may influence the likelihoodthat their usage is passed on via social transmission or
by individual trial-and-error if the behavior occurs commonly enough. This could result in a possible exchange ofmembers and their skills between communities (O’Malley
et al. 2012). However, given the fragmented landscape andpresence of human population across the region, it is notclear that connectivity exists across ape populations (Pielet al. 2013).
Chimpanzees sourced most raw material frombranches close to the ground. However, some sourceswere 2.8 m high, indicating that chimpanzees sometimes climb up to reach desirable sources. Some sourceseven higher up may have not been visible to researchers.
Values for mean height and diameter at which chimpanzees removed raw material at Bilenge are similar to thosereported for chimpanzees at Issa Valley (Almeida-Warren
et al. 2017). This might indicate that chimpanzees focus
on where to source raw material, that is, certain heights or
branch diameters from where bark easily can be detached(Sanz et al. 2004).
Local traditions have been described for animal taxa
as diverse as triggerfish, dolphins, whales and capuchinmonkeys (for review see Fragaszy & Per r y 2003).
However, little is known about the exact time when traditions were invented, for how long they are maintained,
or when they go extinct (Haslam et al. 2016). Thus, as
McGrew (1992; p. 196) succinctly commented about the
chimpanzees of Gombe: “Termite fishing may just as wellhave been invented in 1959, the year before Jane Goodall
arrived, or a million years ago”.
In any case, the current study contributes to theemerging paradigm of “primate archaeology” that aims
to add time-depth to tool use behaviors of extant primate
populations by identifying when and where tool use traditions are invented, modified or go extinct (Haslam et al.
2017).

ACKNOWLEDGEMENTS

Research permission for Mahale Mountains National Parkwas kindly granted by Tanzania National Parks (TANAPA),
Tanzania Wildlife Research Institute (TAWIRI), and TanzanianCommission for Science and Technology (COSTECH). The
author is also thankful to: field assistance by Shaka andSinga Kabugonga; logistical support from Mahale Wildlife
Conservation Society; plant identifications by Frank Mbago,
Botany Department, University of Dar es Salaam; termite identification by Rudolf Scheffrahn, University of Florida, USA.
Special thanks to Edward Kohi, Pius Kavana and Maria Mbogo
for keeping my field equipment safe in Kigoma town; to Michio
Nakamura for help with logistics in the field; to Volker Sommer
for helpful comments on this manuscript; and to reviewer
William McGrew for his constructive feedback. Financial
support was provided by a Leverhulme Trust Early Career
Fellowship and The Boise Trust Fund, University of Oxford.

REFERENCES

Almeida-Warren K, Sommer V, Piel AK, Pascual-Garrido A
2017. Raw material procurement for termite fishing tools
by wild chimpanzees in the Issa valley, western Tanzania.
Am J Phys Anthropol 164:292–304.

Pan Africa News, 24(2), December 2017https://doi.org/10.1002/ajpa.23269

Collins DA, McGrew WC 1988. Habitats of 3 groups ofchimpanzees (Pan troglodytes) in western Tanzania compared. J Hum Evol 17:553–574.
https://doi.org/10.1016/0047-2484(88)90084-X

Fragaszy DM, Perry S (eds) 2003. The Biology of Traditions:
Models and Evidence. Cambridge University Press,
Cambridge, UK.
Goodall J 1964. Tool using and aimed throwing in a community of free-living chimpanzees. Nature 201:1264–1266.
https://doi.org/10.1038/2011264a0

19
Am Nat 164:567–581.

https://doi.org/10.1086/424803

Teleki G 1974. Chimpanzee subsistence technology: Materials and skills. J Hum Evol 3:575–594.
https://doi.org/10.1016/0047-2484(74)90018-9

Uehara S 1982. Seasonal changes in the techniques employedby wild chimpanzees in the Mahale Mountains, Tanzania,
to feed on termites (Pseudocanthotermes spiniger). Folia
Primatol 37:44–76.
https://doi.org/10.1159/000156020

Haslam M, Hernandez-Aguilar A, Arroyo A et al. 2017.
Primate archaeology evolves. Nat Ecol Evol 1:1431–1437.
https://doi.org/10.1038/s41559-017-0286-4

Haslam M, Luncz L, Pascual-Gar rido A et al. 2016.
Archaeological excavation of wild macaque stone tools. J
Hum Evol 96:134–138.
https://doi.org/10.1016/j.jhevol.2016.05.002

Hobaiter C, Poisot T, Zuberbühler K, Hoppitt W, Gruber T
2014. Social network analysis shows direct evidence forsocial transmission of tool use in wild chimpanzees. PLoS
Biol 12:e1001960.

<NOTE>An Old Female Bonobo Carried a
Dead Red-Tailed Monkey for overa Monthhttps://doi.org/10.1371/journal.pbio.1001960

Kazuya Toda1, Nahoko Tokuyama2,& Takeshi Furuichi1

https://doi.org/10.1016/S0047-2484(82)80032-8

1 Primate Research Institute, Kyoto University, Japan
2 The Graduate University for Advanced Studies, Japan(E-mail: toda.kazuya.78x@st.kyoto-u.ac.jp)

McBeath NM, McGrew WC 1982. Tools used by wild chimpanzees to obtain termites at Mt Assirik, Senegal: The
influence of habitat. J Hum Evol 11:65–72.
McG rew WC 1992. Chimpanzee Material Culture:
Implications for Human Evolution. Cambridge University
Press, Cambridge, UK.
McGrew WC, Collins, DA 1985. Tool use by wild chimpanzees (Pan troglodytes) to obtain termites (Macrotermes
herus) in the Mahale Mountains, Tanzania. Am J Primatol
9:47–62.
https://doi.org/10.1002/ajp.1350090106

Musgrave S, Morgan D, Lonsdorf E, Mundry R, Sanz C
2016. Tool transfers are a form of teaching among chimpanzees. Sci Rep 6:34783.
https://doi.org/10.1038/srep34783

Nakamura M, Corp N, Fujimoto M et al. 2013. Ranging behavior of Mahale chimpanzees: A 16 year study. Primates
54:171–182.
https://doi.org/10.1007/s10329-012-0337-z

Nishida T 1968. The social group of wild chimpanzees in the
Mahali mountains. Primates 9:167–224.
https://doi.org/10.1007/BF01730971

Nishida T, Uehara S 1980. Chimpanzees, tools and termites: Another example from Tanzania. Curr Anthropol
5:671–672.
https://doi.org/10.1086/202545

O’Malley RC, Wallauer W, Murray CM, Goodall J 2012. The
appearance and spread of ant fishing among the Kasekela
chimpanzees at Gombe. Curr Anthropol 53:650–663.
https://doi.org/10.1086/666943

Pascual-Garrido A 2017. Termite fishing tools of wild chimpanzees and source plant scars: Towards an archaeology ofthe perishable. Manuscript submitted for publication.
Pascual-Garrido A, Buba U, Nodza G, Sommer V 2012.
Obtaining raw material: Plants as tool sources for Nigerian
chimpanzees. Folia Primatol 83:24–44.
https://doi.org/10.1159/000338898

Piel AK, Stewart FA, Pintea L et al. 2013. The Malagarasi
river does not form an absolute barrier to chimpanzee
movement in Western Tanzania. PLoS One 8:e58965.
https://doi.org/10.1371/journal.pone.0058965

Sanz CM, Deblauwe I, Tagg N, Morgan DB 2014. Insect preycharacteristics affecting regional variation in chimpanzee
tool use. J Hum Evol 71:28–37.
https://doi.org/10.1016/j.jhevol.2013.07.017

Sanz CM, Morgan DB 2007. Chimpanzee tool technology in
the Goualougo Triangle, Republic of Congo. J Hum Evol
52:420–433.
https://doi.org/10.1016/j.jhevol.2006.11.001

Sanz CM, Morgan DB, Gulick S 2004. New insights into
chimpanzees, tools and termites from the Congo Basin.

INTRODUCTION

Carrying infants is one of the most important maternal tasks determining the survival of dependent infants
in non-human primates (Nicolson 1987; Ross 2001). Even
after the death of their infants, mothers tend to carry thecarcasses for several days, and even up to several months(Hosaka et al. 2000; Sugiyama et al. 2009; Tokuyama etal. 2017). Such prolonged carrying can be considered as“maternal persistency” (Sugiyama et al. 2009; Biro et al.
2010).
Excluding mothers that carry their infants, nonhuman primates rarely carry objects for a long period,(Ross 2001; Carvalho et al. 2012). However, at Wamba in
Democratic Republic of the Congo, we observed that an
old female bonobo (Pan paniscus) carried the carcass ofa red-tailed monkey (Cercopithecus ascanius) for over amonth. We here report occasional observations of her behaviors and conditions of the monkey carcass, and discuss
why she had been persistent with the dead body of anotherprimate.

SITE AND SUBJECT

Observations were made in 2016 at Wamba, Luo
Scientific Reserve, Democratic Republic of Congo, which
is a long-term field site for the study of wild bonobos (Kano
1992). There are two adjacent groups of bonobos, the eastern group (PE) and the western group (PW), of which all
members were fully habituated and identified. We usuallyfollowed the PE group from one night’s bed sites (around
06:00 h) to the next night’s bed sites (around 17:00 h), andoccasionally observed the PW group especially during
inter-group encounters. The adult female reported in thepresent study, Chio (estimated to be 45–50 years old), hadbeen identified in the PW group together with her daughter (Chacha: estimated to be 7–8 years old) in September
2012. Since then, Chio had not been observed to have an
infant.

Pan Africa News, 24(2), December 201720

Figure 1. Chio resting on a tree with a mummified red-tailed monkey.

OBSERVATIONSOn August 18, we found that Chio was carrying thedead body of a red-tailed monkey at 13:47 h. The monkey
carcass seemed to be an adult based on size, which was
as large as infant bonobos at one or two years of age. It
did not have any visible external injury and retained its
head, all limbs, tail, and fur. It was suspected to have diedseveral days before the observation because it had alreadydecayed and emitted a decomposing odor to the observers. When Chio rested with the carcass on a tree at 14:00 h,
two adult males approached her to watch it but did not try
to take it. At 14:34 h, Chio hung it on her neck like a muffler and moved away with it.
On August 25, we found that Chio was with the monkey carcass at 13:40 h, but we could observe her only once
on that day.
On August 29, we found that Chio walked on the
ground with the carcass hanging on her neck at 12:46
h. The decay of the carcass was proceeding. Little fur
remained and it was dried up from the head to the tail,including all the limbs. Chio grasped the carcass in her
hand and engaged in genito-genital rubbing with another
adult female at 12:48 h, then she moved away hanging it
on her neck.
On August 31, we found Chio resting on a tree withthe carcass hung around her neck at 06:43 h. One infant
approached Chio apparently to play with the carcass at
07:13 h and Chio moved the carcass from her neck onto
her abdominal region to prevent the infant from touching
the carcass. At 07:21 h after the infant left, Chio lifted thehead of the carcass to eye level and gazed at it for about
five minutes. She groomed it for two minutes at 07:36 h,occasionally flapping flies gathering around it, and moved
away with it at 07:40 h.
On September 17, we found that Chio had the mummified monkey at 1649 h, but we only caught a glimpse ofher once before members made night beds.
On September 21, we found Chio was still in possession of the carcass at 07:14 h. The monkey carcass was

fully mummified. When she engaged in genito-genital
rubbing with another adult female and a juvenile female
at 07:34 h, she grasped the mummy in her hand. Then,
she rested and held the mummy in her abdominal region
at 07:40 h (Figure 1, also see Video 1 available online at
http://mahale.main.jp/PAN/24_2/24(2)_04.html). One infant approached her and watched the mummy at 07:56 h.
She suddenly grabbed the arm of the infant, and the infant
ran away screaming. Chio groomed the mummy with her
mouth at 08:49 h, and moved away with it hanging on herneck.
On September 29 and 30, we found that Chio stillwalked with the mummy hanging around her neck at
11:23h and13:40h. The mummy was intact until our last
observation.
On October 25, we found Chio within members of thePW group at 10:18 h, but she was no longer carrying themonkey carcass.

DISCUSSION

Chio had carried the dead red-tailed monkey for atleast 43 days without a motivation of preying or playing,although we did not observe how she got and released
it. She was observed to groom the hair of the monkey
carcass, gaze at its body, and defend it from curious juveniles. She seemed to handle the monkey carcass carefully
and protectively even during it had been decaying. Her
behaviors towards the decaying monkey carcass could beconsidered similar to the behaviors of mothers carrying
their dead infants with great care (Kano 1992; Hosaka
et al. 2000; Sugiyama et al. 2009; Biro et al. 2010;
Tokuyama et al. 2017).
Few observations of non-human primates carrying
bodies of other animal species for a long time have beenreported until date. In chimpanzees, however, a barren female carried a limb of dead dried red colobus (Procolobus
rufomitratus) for seven hours at Mahale (Hosaka et al.
2000), and an adolescent female carried a dead hyrax(Dendrohyrax dorsalis) for fifteen hours, slept with it in
her nest, and groomed it at Bossou (Hirata et al. 2001).

Pan Africa News, 24(2), December 2017
These observations might indicate that their imagination
could make an unusual perception and reaction towards
other animal species.
Allomothering, defined as temporary carrying of
non-kin infants is performed mostly by non-parous young
females, which has been suggested to be a process of acquiring maternal skills in non-human primates (McKenna
1979). However, it is occasionally conducted by adultparous females without their own infants (Nozaki 2009;
Tokuyama 2015). Chio’s case fits the latter well, because
she had not cared her own infants for a long time. The
unusual carrying might have been a coincidence inducedby the maternal drive for infant caring.

ACKNOWLEDGMENTS

We sincerely thank the Ministry of Scientific Researchof the Democratic Republic of Congo for granting permission to conduct research on bonobos and the Research Centeron Ecology and Forestry (CREF) in the Democratic Republicof Congo for supporting our research in the Luo Scientific
Reserve. We sincerely thank Dr. Sakamaki for his great
contribution to continuous observations and camp management at Wamba and the local assistants for their helps of ourobservations. Field research of KT was financially supported
by Leading Program of Primatology and Wildlife Science(PWS) of Kyoto University and Grant-in-Aid for JSPS Fellows(17J01336), Japan.

21
https://doi.org/2433/203112

Tokuyama N, Moore DL, Graham KE, Lokasola A, Furuichi
T 2017. Cases of maternal cannibalism in wild bonobos(Pan paniscus) from two different field sites, Wamba andKokolopori, Democratic Republic of the Congo. Primates
58:7–12.
https://doi.org/10.1007/s10329-016-0582-7<BOOK REVIEW>

Cannibalism: A Perfectly Natural
History
By Bill Schutt
Chapel Hill, NC: Algonquin Books, 332 pp.
ISBN: 978-1-61620-462-4, 2017. $26.95 (USA), paperback

William C. McGrew
School of Psycholog y & Neuroscience, Universit y of St
Andrews, Scotland, UK(E-mail: wcm2@st-and.ac.uk)

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https://doi.org/10.1007/s10329-008-0118-x

Ross C 2001. Park or ride?: Evolution of infant carrying inprimates. Int J Primatol 22:749–771.
https://doi.org/10.1023/A:1012065332758

Sugiyama Y, Kurita T, Matsui T, Kimoto S, Shimomura
T 2009. Carrying of dead infants by Japanese macaque(Macaca fuscata) mothers. Anthropol Sci 117:113–119.
https://doi.org/10.1537/ase.080919

Tokuyama N 2015. A case of infant carrying against themother’s will by an old adult female bonobo at Wamba,
Democratic Republic of Congo. Pan Afr News 22:15–17.

Given that chimpanzees have shown more instancesof cannibalism than all other species of non-human primates combined, and also that chimpanzees are the mostinvoked species for modelling the evolutionary origins
of human behaviour, this book seemed worth scrutinising. Furthermore, the sub-title suggests a natural science
viewpoint, rather than the more usual social science (i.e.,anthropology) viewpoint, as does the author’s status
as a vertebrate zoologist (on the faculty of Long Island
University, USA). For comparative contrast, see what
may be the best previous example of tackling the topic,
Petrinovich’s (2000) The Cannibal Within, in which nonhuman cannibalism gets only 5 pp.
First, the structure and format. Schutt does not do
research on cannibalism but instead summarises andsynthesises findings on the topic. But he goes beyond
reading and analysing published material by travelling tointerview researchers in situ, with participant observationthat entails (e.g.) archaeology and anthropophagy. (Yes,he eats human tissue.) About this, he writes accessibly but
academically (e.g., providing scientific names for taxa),
with a light touch enlivened by humour. Good points are:28 pages of endnotes by text page number, which provide
references to publications; an extensive book list (n = 68),

22
of which a third focus on cannibalism; 35 line drawings
as illustrations (although almost all lack captions). Not
so good is the inexplicable lack of an index. The result isaward-winning, and the content usefully extends and updates Petrinovich’s older efforts.
What are the book’s strong points? His is the mostcomprehensive, available treatment of cannibalism interms of variation across animal taxa, from invertebrates(cricket, mantis, spider, snail) to vertebrates (spadefoottoad, hamster, polar bear, caecilian, dinosaur). Some ofthis amounts to myth-busting. Among extinct hominins,
he concentrates on Neanderthals, as they provide the best
data. But in all cases, he focusses healthy scepticism on
exaggerated claims, showing how hard it is methodologically to ascertain cannibalism unless observed directly.
He covers the standard sub-categories of types, that is,survival, ritual, nutritional, endo-, exo-, etc., but usefully
adds others that are less well-known, such as filial, medical, auto-, etc.
Schutt includes the standard case studies: New
Guinean mortuary cannibalism by the Fore and the resulting disease of kuru, Donner Party (American settlers
snowbound and starving), Christian Eucharist, etc. In
each case, he is able to go beyond Petrinovich, showingthat research continues to yield new insights. But he shies
away from the Aztecs, adopting an anti-conquistadore,
pro-native American stance that is almost polemical. He
details other cases, e.g., WWII Stalingrad, 20th century
China, and European medical anthropophagy that surprisingly flourished and endured until recently. His coverageof present-day, fashionable placentophagia in the West isentertaining.
What about weak points? The shadow of William
Arens hangs over the book, long after his sell-by date.
(Arens wrote a provocative book, The Man-Eating Myth,almost 40 years ago that claimed that human cannibalism did not exist.) Although he was quickly debunked,
somehow his influence persists, and Schutt spends far
too much time in musing conversations with Arens, whotries to weasel out of the facts. Especially regretable in
Schutt’s ignoring of Anasazi cannibalism in the American
Southwest, which is well-documented archaeologically. At
times, he wanders off-topic, such as telling us more than
we need to know about prions versus viruses. Some of his
forays into the arts come across as a bit trivial (Odysseus,
Robinson Crusoe), while other obvious candidates are ignored, such as a recent, long-running American televisionseries, ‘Dexter’, which features an anthropophagic forensic technician.
But what about non-human primates? Disappointingly, although chimpanzees get most of the mentions, all
but one 3-page section are superficial, and in it, only threesources are cited: Hiraiwa-Hasegawa (1992) (also usedby Petrinovich), Goodall (1977), and a popular reference
to Townsend et al. (2007). This will disappoint readersof PAN, considering all that has been published between
Bygott (1972) and Nishie & Nakamura (2018). All ofthese papers focus on infanticide, while Schutt seems
unaware of (e.g.) consumption of adults in other primate
taxa (Hamalainer 2012) or non-infanticidal consumption(Dellatore et al. 2009).

Pan Africa News, 24(2), December 2017
So, a panthropologist will learn nothing about primate
cannibalism, but the book will put it into comparative perspective, especially if read sequentially after Petrinovich.

REFERENCES

Arens W 1979. The Man-Eating Myth. Oxford University
Press, Oxford.
Bygott JD 1972. Cannibalism among wild chimpanzees.
Nature 238:410–411.
https://doi.org/10.1038/238410a0

Dellatore DF 2009. Two cases of mother-infant cannibalism
in orangutans. Primates 50:277–281.
https://doi.org/10.1007/s10329-009-0142-5

Goodall J 1977. Infant killing and cannibalism in free-living
chimpanzees. Folia Primatol 28:259–282.
https://doi.org/10.1159/000155817

Hamalainer A 2012. A case of adult cannibalism in thegray mouse lemur, Microcebus murinus. Am J Primatol
74:783–787.
https://doi.org/10.1002/ajp.22034

Hiraiwa-Hasegawa M 1992. Cannibalism among non-humanprimates. In: Cannibalism: Ecology and Evolution among
Diverse Taxa. Elgar M, Crespi B (eds), Oxford University
Press, New York, pp. 323–338.
Nishie H, Nakamura M 2018. A newborn infant chimpanzee snatched and cannibalized immediately after birth:
Implications for “maternity leave” in wild chimpanzee. Am
J Phys Anthropol 165:194–199.
https://doi.org/10.1002/ajpa.23327

Petrinovich L 2000. The Cannibal Within. Aldine de Gruyter,
New York.
Townsend SW, Slocombe KE, Thompson ME, Zuberbühler
K 2007. Female-led infanticide in wild chimpanzees. Curr
Biol 17:R355–R356.
https://doi.org/10.1016/j.cub.2007.03.020

Pan Africa News, Vol. 24, No.2
Published in December, 2017
Address: c/o Human Evolution Studies,
Dept. of Zoology, Faculty of Science,
Kyoto Univ., Kyoto, 606-8502, JAPANTEL: (+81)75-753-4108
FAX: (+81)75-753-4115
E-mail: pan.editor@gmail.com
URL: http://mahale.main.jp/PAN/
ISSN: 1884-751X (Print), 1884-7528 (Online)