-finge forest, and survey on the rarest chimpanzee...
TRANSCRIPT
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“Habitat Mapping of the Babanki-Finge Forest, and Survey on
the Rarest Chimpanzee (Pan troglodytes ellioti) in the Bamenda
Highlands, North-West Cameroon”
By Osiris A. DOUMBE
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« I want my children and the future generations to see the chimps. In the past, there were
buffaloes here and there… They all left because of deforestation. Children can only see
buffaloes in pictures now, and they don‟t understand. »
- ALUKEH “Pa” Moses Amah
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Dedication
I dedicate this long and hard work to the memory of my Dad without who I wouldn‟t be half
of the man I am today. This is the first step, the first of many, to be a part of the awareness,
the development, and the emancipation of the Cameroonian people. This is not to prove to
you what I am capable of, or to prove anything to anyone, just because it is the right thing to
do, as the son you educated to love and to be endlessly proud of his fatherland.
Always With Love
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Abstract
The Nigeria-Cameroon chimpanzee (Pan troglodytes ellioti) is the rarest and most endangered
subspecies of chimpanzees. Restricted to the lowland and mountainous forests of Eastern
Nigeria and Western Cameroon, between 3,500 and 9,000 individuals remain in one of the
most populated areas of Africa.
Situated in the heart of the distribution range of Pan troglodytes ellioti, the Babanki-Finge
forest is one of the last remaining montane forests of the Bamenda Highlands (North-West
region of Cameroon). The first purpose of this study was to create the first map of the forest
of Babanki-Finge. Divided in three main ensembles of patches (Alegafor, Abong-Phen and
Kefem) the forest cover totalizes an area of about 400ha. The forest, now concentrated along
watercourses, appears to have undergone much pressure from farmers, hunters and Mbororo
(Fulani) pastoralists. The anthropomorphic pressure is unevenly spread amongst Babanki-
Finge, which results in an unequal distribution of wildlife throughout the different patches.
The most Southern pocket of forest (Alegafor) is highly disturbed by human activities, and
seems empty of any diurnal primates. On the other hand, 71 chimpanzee nests were found in
Abong-Phen and Kefem.
The collected data on the nesting ecology of the Pan troglodytes ellioti of Babanki-Finge, is
the first study on the chimpanzees of the Bamenda Highlands. Generally, the chimpanzees of
Babanki-Finge tend to build nests at a height of about 9m, and to sleep in groups smaller than
three individuals. The results show they prefer building nests on Carapa grandiflora and
Strombosia sp. trees. The short duration of the data collection, and the unreliability of nest
decay rates prevented us from making any measurements regarding the ape density in
Babanki-Finge. The population was estimated to be small and not viable in the long-term. In
addition to chimpanzees, two species of monkeys occur in the North of Babanki-Finge: putty-
nosed monkeys (Cercopithecus nictitans martini) and tantalus monkeys (Chlorocebus
tantalus tantalus).
The creation of forest corridors linking Babanki-Finge to largest forest areas such as Kilum-
Ijim in the North-East and Kom-Wum Forest Reserve in the North-West is indispensable for
the exchange of genes between isolated populations of chimpanzees and the survival of
mammals in the long-term.
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Résumé
Le chimpanzé du Nigéria-Cameroun (Pan troglodytes ellioti) est la plus rare des sous-espèces
de chimpanzés, et la plus en danger. Entre 3 500 et 9 000 individus vivent désormais à l‟Est
du Nigéria et à l‟Ouest du Cameroun, dans une des régions les plus peuplées d‟Afrique.
Située au cœur de l‟aire de distribution de Pan troglodytes ellioti, la forêt de Babanki-Finge
est l‟une des dernières forêts d‟altitude des montagnes de Bamenda (région du Nord-Ouest du
Cameroun). Cette étude a servie, dans un premier temps, à établir la première carte de la forêt
de Babanki-Finge. Divisée en trois ensembles (Alegafor, Abong-Phen et Kefem) la couverture
forestière totalise environ 400ha. Concentrées dans les vallées de rivières, les poches de forêt
semblent avoir subies beaucoup de pression de la part des fermiers, des chasseurs, et des
pasteurs Mbororo (Peuls-Foulbés). La pression anthropomorphique est inégalement répartie à
travers la forêt, d‟où une distribution disparate de la faune sur l‟ensemble des fragments :
l‟ensemble forestier le plus au Sud (Alegafor) est très perturbé par les activités humaines, et
aucun singe ne semble la visiter, alors que 71 nids de chimpanzés ont été trouvés à Abong-
Phen et Kefem.
La seconde partie de cette thèse, centrée sur l‟écologie de la nidification de Pan troglodytes
ellioti de Babanki-Finge, est la première étude sur le chimpanzé des montagnes du Nord-
Ouest Cameroun. Généralement, les chimpanzés de Babanki-Finge construisent des nids dans
des arbres, à environ 9m du sol, et dorment en petits groupes de moins de trois individus. Les
données relevées montrent qu‟ils préfèrent construire des nids dans des arbres appartenant aux
espèces Carapa grandiflora et Strombosia sp. La courte durée du travail sur terrain et le
manque de fiabilité du taux de décomposition des nids nous a empêché de faire toute
estimation sur la densité des chimpanzés de Babanki-Finge. La population a été estimée
comme étant petite et non viable sur le long terme. Deux autres espèces de singe ont
également été observées dans le Nord de Babanki-Finge : le singe hocheur (Cercopithecus
nictitans martini) et le singe vert (Chlorocebus tantalus tantalus).
La création de corridors de forêts rattachant Babanki-Finge à de plus larges forêts telles que
Kilum-Ijim au Nord-Est et la Réserve Forestière de Kom-Wum au Nord-Ouest, est
indispensable pour permettre les échanges génétiques entre des populations de chimpanzés, et
la survie des mammifères à long terme.
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Acknowledgements
I have to start by her, it‟s mandatory… I want to first give thanks to my wonderful mother, for
her constant support, her inconditional love, and her trust in me. You are the best Maman,
merci pour tout!
I will be forever grateful to Le Conservatoire de la Vallée des Singes. Thanks to their grant, I
was allowed to accomplish my field work in Cameroon. Without them, I could definitely not
have gone that far in my research. Thank you VERY much.
I particularly want to thank my tutor Dr Caroline Ross for her help and advices regarding my
thesis. I want to extend my gratitude to Dr Mary MacKenzy who has always been there for
me. Without her knowledge, I wouldn‟t have come up with such beautiful maps, that‟s for
sure! Thank you once again.
In extension, I would like to thank all the professors of the MRes program of Primate
Biology, Behaviour and Conservation. Their teaching and advises throughout the year surely
helped me in the writing of this final document. I also want to thank Pr. Sarah Gartland for
her advices in English writing, thank you.
Special thanks to Dr Jean-Luc Mouget for his kindness and advices.
Merci beaucoup à toute ma petite famille adorée! Herna, Liko, Sanga et Neph‟, merci à vous
pour votre amour, pour être toujours présents pour moi, et soudés quand je suis loin. Bisous !
J‟aimerais également étendre ma gratitude à Papi et Mamie pour les petites attentions, à Tata
Suzy et Tonton Emile pour leurs emails dans la brousse, et au reste de ma grande famille pour
leur soutien. Merci à tous ! Much Love.
None of this would have been feasible without the help and support from the local team in
Bamenda and Babanki-Finge. I would like to show my deepest appreciation to John DeMarco
who believed in this project and did everything he could to help me. I would like to offer my
special thanks to my field assistants Ernest, David (and his Madam Atchu!), and, last but not
the least, Pa‟ Moses (I do miss your wife‟s grilled sweet potatoes!). All of your work during
my time in the North-West Cameroon was deeply appreciated, thank you once again.
In addition, I want to the Kedjom-Keku NGO and particularly Bara for her welcoming, and
Robert for his help.
This field work wouldn‟t have been much more than work if it wasn‟t for the rest of my
Cameroonian family. I want to particularly thank Mama, Uncle Talla, Esther (Esta!), Sama,
and the lovely kids (nope, don‟t call me Uncle White Man, it‟s Uncle Osi) Josh, Aissatou, K
and John for their love. Thank you also to Mama‟s sister, Mbong and Antie Lau in Yaoundé.
My thoughts also go to my big brothers Gérard and Wally who picked me up at the airport at
03.00am in Douala. Much love and appreciation also go to my brother Eric from Bamenda.
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I want to thank my British Family: Sally (British Mum), Roy, Rita & Jim (thank you for the
survival kit) and the whole neighbourhood, much love to Morden! In extension I also want to
thank Fanni, Godefroy and my man Joao! Speaking of which, I would like to thank very much
my classmates: Shai, Simon, Ivette, Rachel, Flic, Samara, Piotr, Rosemarie, Melissa and the
tarsier girl Sally.
No, no Mum Lashell, I‟m not forgetting you. Thank you very much for your help. Thanks
also to Sasha for supporting my work and for... Well, just for being here, ha!
Ofir… Where should I start? What a pleasure and an inspiration to finally meet the person
behind LAGA after several years reading and watching interviews about. Thank you so much
for the support, the documents, and the websites which helped me to build a much better-
constructed conservationist discussion, and gave me a new perception on the problems of
animal trafficking in tropical Africa. I also would like to extend my gratitude to Luc Mathot.
Thank you to Arthur who always had some time for me, even in the wildest places of
Cameroon and Chad. Hopefully I will have the pleasure to meet you in Central Africa in the
close future.
Merci également au Docteur Bakwo Fils Eric Moïse de Maroua pour sa spontanéité et sa
coopération.
I would like to thank deeply Guy, Adrienne, Danika, Jessica and Rahina and their whole
family and friends, for their presence in the end of my stay in Cameroon.
Finally, thank you to Billy the chimp and his two friends locked in a cage somewhere in the
countryside of North-West Cameroon. Thank you, you have brought the words of this
dissertation closer to the truth and essential the improvement of the conservation of the
Cameroonian wildlife. I truly hope you‟ll be soon placed in a sanctuary…
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Contents
Chapter 1 – Introduction 1
1 The Cameroon Highlands 1 1.1 Description 1
1.1.1 Geography & History 1
1.1.2. Plant and Animal Endemism 3
1.2 Anthropomorphic Pressure 5
1.2.1 Human Migrations and High Densities 5
1.2.2 Bushmeat and Wildlife Medicine 6
1.2.3 Management of Resources 9
2 The Chimpanzee 10 2.1 The Genus Pan 10
2.2 Pan troglodytes 11
2.2.1 How many Subspecies? 11
2.2.2 Behaviour and Ecology 15
2.2.3 Conservation Status 16
2.3 Pan troglodytes ellioti 17
2.3.1 The Most Threatened Subspecies of Chimpanzees 17
2.3.2 “The Great Apes of Tubah” 18
3 Estimating the Population of Chimpanzees 19
4 The Study 21 4.1 The Montane Forests of Tubah Sub-Division 21
4.2 Aims & Objectives 22
Chapter 2 - Methods 23
1 Study Site 23
2 Data Collection 24 2.1 Study Period 24
2.2 The Global Positioning System (GPS) 26
2.3 Forest Delimitation 26
2.4 Diurnal Primate Signs 27
2.4.1 Direct Signs 27
2.4.2 Indirect Signs 27
Indirect Signs – Nest Recording 27
Indirect Signs – Other Signs 29
2.5 Habitat Description 29
2.6 Fig Tress 33
3 Data Analyses 33 3.1 Creating a Map 33
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3.2 Analysing Recorded Signs 33
3.2.1 Decay Rate 34
3.2.2 Nest Characteristics 36
Chapter 3 – Results 36
1 Map of Babanki-Finge 36 1.1 Alegafor 36
1.2 Abong-Phen 38
1.3 Kefem 42
1.4 Comparisons Between Areas 43
2 Observations and Signs of Monkeys and Apes 45 2.1 Direct Signs 45
2.2 Indirect Signs 46
2.2.1 Chimpanzee Nests
Chimpanzee Nests – Tree Species 49
Chimpanzee Nests - Height 52
Chimpanzee Nests – Cluster Size 53
Chimpanzee Nests – Nesting Site Choice 54
2.2.2 Other Signs 55
Chapter 4 - Discussion 55
1 Forest Mapping 55 1.1 Characteristics of the Vegetation 55
1.1.1 Forests 57
1.1.2 Aframomum sp. 57
1.1.3 Grassland 56
1.2 Human Impact 59
2 Pan troglodytes ellioti: Discussion and Comparisons 60 2.1 Chimpanzee Density 61
2.1.1 Nest Counting 62
Accuracy of Decay Rate 62
2.1.2 Nest Density 65
2.2 Nest Height 65
Terrestrial Nests 66
2.3 Cluster Sizes 69
2.4 Tree Species 72
2.5 Nesting Site Choice 73
3 Other Wildlife 73 3.1 Monkeys 73
3.2 Mammals and Birds 75
4 Comparing Babanki-Finge with Other Fragmented Forests 76
5 For a Better Conservation 77
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5.1 Hopes for Babanki-Finge (?) 78
5.1.1 Touristic Potential 79
Touristic Potential – Bird Watching 80
Touristic Potential – Site-Seeing 80
Touristic Potential – Primate Tourism 82
5.2 Conservation in the North-West Region of Cameroon 83
6 Future Research 85 6.1 Babanki-Finge 85
6.2 The North-West Region of Cameroon 87
6.3 Recommendations 87
7 General Conclusion 88
References 90
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List of Figures and Tables
Figures
Figure 1: Map of the Cameroon line and the distribution of the Cameroonian forest 2
Figure 2: Distribution of Chimpanzees and Bonobos 14
Figure 3: Satellite map of the Forest surrounding the villages of Kedjom-Keku and
Kedjom-Ketingo (Babanki), Bambui and Finge 21
Figure 4: Satellite map of the Babanki-Finge forest 24
Figure 5: Drawing representing the requisite measurements for the assessment of the
height of the chimpanzee nest 28
Figure 6: Photograph of a mature Cyathea manniana 32
Figure 7: Photograph of an unknown species of trees 32
Figure 8: Satellite Photograph of a part of Alegafor with waypoints 34
Figure 9: Map of Babanki-Finge and the towns surrounding the forest 37
Figure 10: Photograph of the middle part of Alegafor 38
Figure 11: Map of Alegafor and the Planted Acacia Forest 39
Figure 12: Grassland in Southern Ndong-Efuh 40
Figure 13: Plantation of djama-djama and maize on the edge of the sub-patch of
Ndong-Efuh 40
Figure 14: Map of Abong-Phen 41
Figure 15: Photograph of Abong-Phen 42
Figure 16: Map of Kefem 44
Figure 17: Old nest of Chimpanzee in the canopy 47
Figure 18: Map of the chimpanzee nesting sites in Abong-Phen 50
Figure 19: Map of the chimpanzee nesting sites in Kefem 51
Figure 20: Exploded pie representing the species of trees used by the chimpanzees of
Babanki-Finge for nesting 52
Figure 21: Distribution of the chimpanzee nests in the layers of the trees of the forest 53
Figure 22: Frequency of chimpanzee nest group sizes in Babanki-Finge 54
Figure 23: Map of the forest patches of Babanki-Finge 56
Figure 24: Photograph of fruits and leaf of Aframomum sp. 58
Figure 25: Map of the sites mentioned in this section 61
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Figure 26: Photograph of a water fall in Fintse 80
Figure 27: Photograph of a water fall in Mendong I 81
Figure 28: Photograph of an Egyptia rousete caught in Ndong-Efuh 81
Figure 29: Photograph of a Noack‟s roundleaf bat caught in Ndong-Efuh 82
Figure 30: Map of the mentioned areas of the North-West region of Cameroon 85
Figure 31: Satellite photograph of the East of Babanki-Finge 86
Tables
Table 1: Descriptive of the estimation of the age of nests 28
Table 2: Vegetation Descriptive 30
Table 3: Sub-categorization of the forest descriptive 31
Table 4: Location, date of first observation, and number of days until the end of the study
for each fresh nest collected 35
Table 5: Cover areas of the different vegetation types encountered in the three ensembles
measured 45
Table 6: Cover of farms and areas with signs of former human sedentary pressure 47
Table 7: Characteristics of the Recorded nests 47
Table 8: Species of trees in which chimpanzee nests were found 52
Table 9: Descriptive Statistics of the measured height of 40 nests 53
Table 10: Number of Nests per Sleeping Site 54
Table 11: Cover areas of the different vegetation types encountered in the three
ensembles measured 60
Table 12: Comparisons of density estimations from previous studies 64
Table 13: Comparison of chimpanzee nest densities between some patches of Babanki-
Finge and other sites 66
Table 14: Comparison of nest heights between Babanki-Finge and populations of Pan
troglodytes spp. And Pan paniscus from other sites 67
Table 15: Comparison of the sizes of sleeping sites between the nest groups of Babanki-
Finge and the ones of Pan troglodytes and Pan paniscus from other sites 70
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Chapter 1 - INTRODUCTION
1. The Cameroon Highlands
1.1. Description
1.1.1. Geography and History
Cameroon is a central African country characterized by ten different phyto-geographic zones
(Olivry, 1986), more than 200 tribes (Onana, 2005), and 48% of Africa‟s known mammals
(Besong, 1995). It is the 4th
richest African country in terms of biodiversity (Eyebe et al.,
2012) and often described as miniature Africa (Molua, 2012; Tchindjang & Fodouop, 2003).
The exceptional diversity of Cameroon is due to both its spread of latitude, and the
mountainous chain which crosses the country in the middle: the Cameroon Line (Olivry,
1986) (Figure 1). This high-altitude horst-like massif is the only known within-plate alkaline
volcanic province (Fitton, 1987). It consists of a chain of old to recent volcanoes (Fitton &
Dunlop, 1985) whose first peak (Mt. Bangou in the South-West region of Cameroon) was
formed during the Eocene (Fosso et al., 2005). This unique 1600km-long mountainous chain
is half oceanic and half continental: it starts in Pagalu (an Equatorial Guinean island), goes
through the Mambila Plateau (in Nigeria, close to the Cameroonian border), and ends in the
Adamawa Mountains (in the Centre of Cameroon), creating an arc between the South-West
and the Centre of Cameroon (Gountie Dedzo et al., 2011; Suh et al., 2001). Several craters,
presumably from the late Pleistocene or the Holocene, are concentrated in this massif
(Tamura, 1990). Furthermore, there are 12 main volcanic centres (Halliday et al., 1990) which
include some of the most imposing peaks of the continent (Figure 1).
Overhanging the Bight of Biafra, Mt. Cameroon, the Bamenda Highlands and the
Clarence Peak of Fernando Pô are the three highest mountain masses of West Africa (Boghey,
1955). Mt. Cameroon is the highest West-African Mountain (peak Fako at 4095m) and one of
the most active volcanoes of the continent, with seven eruptions within the last 100 years (Suh
et al., 2003). On the slope of this volcano, facing the Atlantic Ocean, Debunscha Cape is the
second wettest site in the world (>10,000mm/year) after Cherrapunji in India (Tchindjang &
Fodouop, 2003). From the coastal areas towards the interior of the continent, rainfall
decreases on the successive peaks of the Cameroon line (Molua, 2012): 4,891mm/year on Mt.
Kupé (Bamboutos mountains in the South-West region) (Hofer et al., 2000), 2,100mm/year
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on Mt. Oku (Bamenda Highlands in the North-West region) (Tye, 1992), and finally
1,700mm/year on the Tchabal Mbabo Mountains (Adamawa) (Herrmann et al., 2007).
NIGERIA
Bioko Island
(EQUATORIAL
GUINEA)
100 km
60 miles
S
N
W E
LEGEND :
: Forest
: Mosaic Forest-Savannah
: Cameroon line
: Mountain massifs
Mt. Cameroon (4095m): Major peaks
: Main rivers
Mt. Cameroon
(4095m)
Mt. Oku (3008m)
Mt Tchabal Mbabo
(2460m)
Mt. Manengouba (2411m)
Mt. Kupé
(2064m)
Adamawa
Bamboutos
Mountains
Adamawa
Bamenda
Highlands
2500 km
1500 miles
Figure 1: Map of the Cameroon line and the distribution of the Cameroonian forest. Based on the
Forest Atlas of Cameroon: http://www.wri.org/tools/atlas/map.php?maptheme=cameroon.
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Separated to the Congo Basin by the large Sanaga River in the South of Cameroon, the
lowland and montane forests of Western Cameroon have long been recognized as highly
diverse and have recently been identified as a biodiversity hotspot both continentally and
internationally (Bergl et al., 2006) (Figure 1).
1.1.2. Plant and Animal Endemism
According to paleo-climatology researches, the climate variation within the last 20,000 years
provoked many fluctuations in the entire country but the South-West region which has always
been equatorial humid (Olivry, 1986). In 20,000-18,000 B.C., during the last Ice Age (late
Pleistocene), the expansion of the ice in the Northern and Southern hemispheres provoked the
decreasing of the global sea level which turned to drier conditions on continents, and the
extension of the Sahara desert in Africa (Reynolds, 2005). This climate change reduced
rainfalls up to 30% in some regions (Linder, 2008). During this dry period, South-West
Cameroon (from the Cross River which delimits the current border with Nigeria, to the
Sanaga River), as well as other ecological islands such as the Eastern part of the Democratic
Republic of Congo (DRC), remained one of the few refuges for African tropical plants and
animals (Brenac, 1988; Colyn et al., 1991; Linder, 2008). This dry climate persisted in Africa
until the Holocene (12,000-11,000 B.C), epoch when the ices of the Northern and Southern
hemispheres melted, and the African tropical forest expanded (Brenac, 1988; Reynolds, 2005)
to reach its maximum extension in 8,000-4,500 B.C. (Olivry, 1986). It has been largely
accepted that ecological islands of the Pleistocene glaciations correspond to current centres of
high diversity and endemism (Colyn et al., 1991). Therefore, Western Cameroon became one
of the richest regions in endemic taxa and one of the international biodiversity hotspots of
global significance (Bergl et al., 2006; Tropek & Konvicka, 2010).
The natural habitat of the Cameroonian Highlands is composed of sub-montane and
afromontane forests, with subalpine grasslands on the highest peaks (Fako and Oku) (Ingram
& Nsom Jam, 2007). Sources of the most important streams of the country start in the altitude
of the Cameroon line: in the Western Highlands, for Noun and Cross rivers, and in the
Adamawa Massif for Benoue, Djerem, Lom, and Sanaga rivers (Olivry, 1986). The latter has
played an important role in the local diversity of plants and animals. Having its source in the
savannah of the Adamawa Mountains, this large river separates from North to South the
Western Cameroon forests with the forests of the Congo Basin. The Sanaga River has been
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proposed as a taxonomic barrier (Gagneux et al., 2001; Morgan et al., 2011). Because of this
river, the Western Cameroon forests (along with the Eastern-Nigeria forests) present a
significantly different wildlife from the bigger forest bloc which covers Eastern Cameroon,
Congo, Gabon, Equatorial Guinea, The DRC, Northern Angola (Cabinda), and the South-
West of the Republic of Central Africa. Along the Sanaga River live sympatric closely related
taxa: e.g. drills (Mandrillus leucopheus) and Nigeria-Cameroon chimpanzees (Pan
troglodytes ellioti) live on the North-Western bank of the river, while mandrills (Mandrillus
sphinx) and central chimpanzees (Pan troglodytes troglodytes) inhabit the rainforests of the
South-Eastern bank of the Sanaga (Grubb, 1973; Morgan & Abwe, 2006). Limited by the
Sanaga River in the East and the Niger Delta in the West, the unique wildlife of the Nigeria-
Cameroon trans-boundary region has evolved to combine important levels of endemism and
great richness of species (Blackburn et al., 2010; Morgan et al., 2011).
The Western Highlands of Cameroon possess the largest remaining afromontane forests in
West Africa (Ingram & Nsom Jam, 2007; Tropek & Konvicka, 2010). In terms of plant
species diversity per km², the South-West and North-West provinces of Cameroon are the
richest areas in Africa (Cheek et al., 2001), and the region around Mt. Cameroon has the
highest number of vascular plant species in the continent (Barthlott et al., 1996). This
Ecoregion supports 100 rare/endemic species of plants including taxa with extremely limited
cover; species like Alchemilla fischeri and Newtonia camerunensis are only found on Kilum-
Ijim summits (North-West region) and especially on high altitude in Mount Oku (Ingram &
Nsom Jam, 2007). The Cameroonian Highlands, which support 35 endemic bird species, are
estimated to be the third richest African region for birds (Ingram & Nsom Jam, 2007). The
high altitude of these mountains creates typical environments where several species are
related to taxa from Eastern African mountains such as the Cameroonian subspecies of the
Stuhlmann‟s golden mole (Chrysochloris stuhlmanni balsaci: Lamotte & Petter, 1981) and
mountain reedbucks (Redunca fulvorufula adamauae: Pfeffer, 1962). They also have a high
level of endemism in reptiles (ten endemic species: McKay & Coulthard, 2000; including
seven taxa of chameleons: Gonwouo et al., 2006), amphibians (Zimkus, 2009) and butterflies
(Tropek & Konvicka, 2010). Located in the South-Eastern edge of the Cameroon line, Mt.
Nlonako presents a particular interest for amphibians and reptiles. The reptilian inventory of a
survey conducted between 1998 and 2004 on this mountain, and summarized by Herrmann et
al. (2005a), recorded 89 species including 63 snakes. The authors of this study described this
peak as the richest single-locality area in reptiles in Africa, and probably the richest site in
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snake diversity in the world. Mt. Nlonako has also the richest diversity in amphibians in
Africa (Herrmann et al., 2005b).
The Bamenda Highlands is the most diverse and important area in Western Cameroon after
Mt. Cameroon and Mt. Kupé (Sedláček et al., 2007). Within the Ecoregion of the
Cameroonian Highlands, several taxa are endemic to the Bamenda Highlands (Ingram &
Nsom Jam, 2007; Ndenecho, 2009) and particularly to its highest peak: Mt Oku (Cheek &
Csiba, 2000). These mountains are well-known for their richness in birds (Ndenecho, 2011;
Reif et al., 2007), with several endemic species (Ingram & Nsom Jam, 2007), including the
banded wattle-eye (Platysteira laticincta) and the Bannerman‟s turaco (Tauraco bannermani)
(McKay & Coulthard, 2000), an emblematic bird for local communities but highly localized
and threatened by hunt (Ingram & Nsom Jam, 2007). Several species of primates, including
Nigeria-Cameroon chimpanzees and Preuss‟s guenons (Cercopithecus preussi), (both taxa
considered as Endangered by the International Union for the Conservation of Nature (IUCN),
and endemic to the Bight of Biafra: IUCN, 2013), live in the remaining patches of sub-
montane and afromontane forests of the Bamenda Highlands (Ingram & Nsom Jam, 2007).
However because of the high level of anthropomorphic pressure and the lack of scientific
research in the North-West province of Cameroon, little is known in the distribution and
abundance of primates in the Bamenda Highlands.
1.2. Anthropomorphic Pressure
1.2.1. Human Migrations and High Densities
Burges et al., (2007) have shown human dense areas are positively correlated with species
richness and endemism all over sub-Saharan Africa. The Bamenda Highlands confirm this
statement. More than 1,500,000 people populate this area (Ingram & Nsom Jam, 2007), where
the population increase is about 2.3%/year (Ndenecho, 2009). This region possesses a density
of 100-250 people/km²: it is one of the most populated areas of Cameroon (Morgan et al.,
2011). The North-West region of Cameroon has a long history of human settlements. It is
actually widely accepted that it is the birthplace of the Bantu people (major African group of
tribes which gather a common mother-language), now widespread all over Central, Eastern
and Southern Africa (De Maret et al., 1987). The Bamenda Highlands (commonly called “The
Grassfields”) seem to have been continuously and quite densely inhabited for thousands of
years (Warnier, 1984).
6
More recently, Mbororo cattle grazers (related to the Fulani of West Africa) have moved to
this region, attracted by the grasslands of the mountains. The last wave of migration took
place in the early 1960s, when Bamenda and its periphery were regarded as one of the main
destinations for Bamiléké migrants running away from the rebellion-repression cycle in the
West region organized by France and the newly independent Republic of Cameroon
(Fodouop, 2010). The successive influxes of new communities in the North-West region of
Cameroon have caused an increasing anthropomorphic pressure upon wildlife.
1.2.2. Bushmeat and Wildlife Medicine
Considered as one of the biggest threats to wildlife in tropical forests (Milner-Gulland &
Bennett, 2003) and as the main cause of the decreasing of African ape populations (Pearce &
Ammann, 1995), the commerce of bushmeat is particularly critical in Africa. In the Congo
Basin, between 1 million and 3.4 million tons of wild meat are consumed each year (Wilcox
& Nambu, 2007). This number represents a “boom” of wild animal consumption from which
several scientists predict a crash of the African tropical diversity in a close future (Barnes,
2002).
In West and Central Africa, the amount of primates killed for bushmeat is widely recognized
as unsustainable (Bowen-Jones, 2002). Located at the limit between West and Central Africa,
and sharing taxa with both regions, Cameroon is an important country in wild animal
trafficking and consumption. In his Review of the Commercial Bushmeat Trade on
Central/West Africa, Bowen-Jones (1998) listed Cameroon as the country with the most
references, representing 21% of the literature out of nine countries. The importance of this
traffic in Cameroon has been observed in the legal and illegal markets of the most populated
cities: Douala (the economical capital) and Yaoundé (the political capital). As an example, in
2006, Edderai & Dame conducted a survey in Yaoundé on the bushmeat trade, and discovered
15 markets were selling wildlife products and 145 restaurants/cafeterias were selling 1052
bushmeat dishes per day. The meat, consumed fresh or smoked, is imported via an important
network using both roads and railways. It has been estimated trains unload about one ton of
smoked bushmeat every day in Yaoundé only (Bowen-Jones, 1998). This bushmeat
commercialization is provoked by an ever-growing demand which is due to a dramatically
increasing population together with better infrastructure and communication (Linder & Oates,
2010; Wilcox & Nambu, 2007). Unlike subsistence hunting, commercial hunting is not
7
sustainable for wild animal populations as commercial hunters kill 10 times more animals
than subsistence hunters (Bowen-Jones, 1998). Nevertheless, the main reason for the
important demand of bushmeat, perhaps more than its taste, is its low price. In their study
around the Banyang Mbo Wildlife Sanctuary, in South-Western Cameroon, Wilcox & Nambu
(2007) actually stated the most expensive bushmeat (pangolin, Smutsia spp) was half the price
of the cheapest alternative meat (740FCFA/kg in comparison to 1400FCFA/kg for chicken
meat). It is however important to notice these prices were collected in the country side, where
bushmeat is less expensive than in big cities (Wilkie et al., 2005).
In 1994, a set of laws has been adopted to regulate animal trafficking in Cameroon (Chi,
2007). The management of wild animals has then been reinforced with the creation of the
classes A, B and C, grouping the Cameroonian fauna into three different levels of protection
(Njike, 2007). But despite heavy sentences (the penalty for poaching can be up to a fine of
USD 20,000 and/or an imprisonment for 3 years: Forboseh et al., 2007), for long, wildlife law
has been flouted. To resolve this critical situation, Ofir Drori, a passionate Israeli activist,
created the first law enforcement NGO in 2002: The Last Great Ape Organization (LAGA).
Basing its actions to fight corruption in the dense network of animal trafficking and poaching,
LAGA contributed, since 2006, to one major arrest per week, resulting more than 80% of
these traffickers behind bars (Ofir Drori pers. comm.). Recently, replications of LAGA
throughout Central Africa has extended the fight against corruption and dramatically
improved the number of prosecutions in this region of the continent: the “Projet d‟Appui à
l‟Application de la Loi sur la Faune sauvage” (PALF) created in 2008 in Congo, the
“Renforcement de l‟Application de la Loi Faunique” (RALF) created in 2009 in the Central
African Republic, and the Conservation Justice created in Gabon in 2010.
Wild animal populations are getting scarcer, and therefore protected areas which lack
conservation activities and efficient anti-poaching management become isolated refuges
threatened by increasing hunting (Dupain et al., 2004). The problem of commercial poaching
has now largely entered the limits of protected areas. Some wildlife sanctuaries have actually
been reported to undergo rates of hunting 2 to 13 times higher than sustainable rates (Hughes
et al., 2011). Korup National Park, which was once described as one of the richest and most
diverse rainforests of the world (Korup – An African Rainforest, 1982), has seen its primate
richness drastically falling within the last decades. The first complete study on the impact of
poaching on monkeys in this protected area was made by Linder and Oates (2011). The
results of this survey showed drills were the second most harvested monkeys in the forest in
8
1988 (representing 27% of the carcasses), but the least hunted in 2004-2005 (3.5% of the
carcasses) due to overexploitation. Moreover, no Mandrillus leucophaeus was seen by the
research team in 2004-2005. Out of the seven monkey species studied, only two (the mona
Monkey (Cercopithecus mona) and the putty-nosed Monkey (Cercopithecus nictitans
martini)) were defined as relatively tolerant to hunting. The low density of most large
mammals in Korup National Park has been explained as a consequence of unsustainable
hunting (Bowen-Jones, 1998). Research conducted by Dupain et al. (2004) and Eyebe (2012)
have shown that important populations of gorillas (Gorilla sp.) and chimpanzees occur more
outside than inside protected areas. In Ntonga, a site on the Northern periphery of Dja Faunal
Reserve in South-Eastern Cameroon, the densities of central chimpanzees and Western
Lowland gorillas (Gorilla gorilla gorilla) were found to be respectively 37-55% and 103-
127% higher than inside the faunal reserve borders (Dupain et al., 2004). These numbers raise
questions about the efficiency of national parks, sanctuaries and reserves whose official
statuses and lack of anti-poaching management finances may tend to attract poachers.
Trans-border trafficking with the Republic of Central Africa (RCA), Congo, Equatorial
Guinea, Gabon and Nigeria is another major problem for the protection of the Cameroonian
wildlife. Amongst these body parts, an important portion is used in traditional medicine.
The traffic of primates for medicinal purposes is a worldwide problem and affects many taxa.
Alves et al (2010) showed that 101 species of primates from 38 genera and 10 families were
exploited for wildlife medicine. In the country side of Western Cameroon, people still use
traditional medicine in favour of “western medicine” as it is generally much more affordable
and believed to be more efficient. In their study in the Bechati-Fossimondi-Bessali area in
2011, Etiendem et al. found 62% of the participants suggested that some diseases which could
not be cured by conventiona medicine could be healed by the consumption of gorilla body
parts. Therefore, wildlife medicine adds another significant pressure on animals. Efforts have
been made by local NGOs (especially in the North-West and South-West regions) in the last
years to sensitize people to the problem of the unsustainability of wildlife medicine.
Productions such as the movie “Wildlife Medicine” by AFRICAROOT PRODUCTION
(2012) try to explain to local people the importance of the protection of wild animals toward
their use as remedy. Nevertheless, traditional/ancestral beliefs can also be used in cooperation
with wildlife laws, for some local taboos forbid hunting and eating certain species of animals.
In the Western regions of Cameroon, traditional and religious beliefs are still largely
respected, and local chiefs (Fon) are strongly esteemed (Abbot & Thomas, 2001). The respect
9
of taboos has consequently prevented the extinction of the Cross-River gorillas (Gorilla
gorilla diehli) of Bechati-Fossimondi (Etiendem et al, 2011), and the Nigeria-Cameroon
chimpanzee populations of Mt. Cameroon (Forboseh, 2007), despite being located a few
kilometres only from Douala, the most populous city of Cameroon.
A powerful traditional taboo is however not enough to stop the threat upon wildlife as it is
revealed in Ruth Wiseman‟s thesis (2008) conducted in the Kagwene Gorilla Sanctuary. The
Cross-River gorillas of this protected area located in the North-West of the South-West
provinces of Cameroon have actually been protected by a traditional hunting prohibition. Still,
it has suffered much human pressure, and by 2008, when the area received the official
protection status of sanctuary, 43.26% of the forest of KGS was already denuded in grazing
lands and farms, two environments unsuitable for gorillas.
1.2.3. Management of Resources
The soils of the Cameroonian Highlands, from Mt. Cameroon to the North-West region, are
well-known for being extremely rich in minerals and humus (Olivry, 1986). However, the
high topography and the rocky features have been serious obstacles to the development of the
region until recently (Fongjong, 2008). According to Ndenecho (2009), the agricultural value
of the region has drastically increased in the early 1990s because of road improvement. The
following intensification of farming and the abandon of the traditional fallow resulted in a fast
impoverishment of the soil despite its original richness, as well as a massive deforestation. In
about 40 years, more than 80% of the forest was cleared, reducing the remaining cover to only
3% in 2000 (Ingram & Nsom Jam, 2007). Ndenecho (2009) also commented the continuous
need for new arable soils has provoked the decreasing of grazing lands, forcing cattle and
sheep to invade wild environment, also contributing to the fragmentation of sub-montane and
montane forests. Finally, overgrazing and burning have considerably affected the
geographical features, exposing the soil to erosion. The exploitation of wood for fuel is
another significant threat to the last remaining patches of forest in the Bamenda Highlands
(Ndenecho, 2010a). The high level of anthropomorphic pressure (Tchatchouang et al., 2012)
has provoked a savannization of the tropical montane cloud forests (Ndenecho, 2005) and a
dramatic decreasing of biodiversity. In addition, this region does not possess any protected
area with a status more important than wildlife reserve or wildlife sanctuary.
10
In order to reduce the human pressure on the environment and to practice a more durable
agriculture, environmental NGOs have locally been created since the early 1990s (Ndenecho,
2009). Unfortunately, the life span of these organizations is relatively short in general as they
heavily depend on external funds. Moreover, NGOs in the North-West region tend to have a
short area of action and have therefore a contestable efficiency (pers. obs.). Finally, the last
and probably the most important problem of these organizations, is many of them have been
created, not to produce a change, but for personal enrichment (pers. obs.).
In areas where natural environment still remain, a sustainable management of the use of plants
and animals is urgently needed. Ndenecho (2010b) recorded 86 useful species of plants only
for Mt Oku. The exploitation of these plants for medicine purposes represents a serious threat
for natural habitats and wild animals. This pressure has become even more important since the
international recognition of medicinal values of certain indigenous species. Therefore, Ingram
& Nsawir (2007) argued the overexploitation of Prunus africana (commonly known as „Iron
Wood‟ or locally as „Pygeum‟ or „Kanda Stick‟) for export to Western countries where it is
used to treat benign prostate hyperplasia, has provoked the extermination of close to 50% of
the mature trees in the North-West region of Cameroon.
2. The Chimpanzee
2.1. The Genus Pan
In his publication on the genetic similarities between great apes and Humans (Homo sapiens),
Jonathan Marks (2002) argued the scientific classification of chimpanzees has been hazardous
because of their resemblances with Homo sapiens. Published in 1758, the tenth edition of
Systema naturae (System of Nature) by Carl Linnaeus is recognized as the foundation of the
modern biological classification. The Swedish botanist and physician found the case of the
chimpanzee so confusing, that he decided to divide the reports of the species in two
classifications: Homo troglodytes (also named Homo nocturnus) belonging to the same genus
as Humans, and Simia satyrus belonging to a separate primate genus. Several biologists
criticized this ambiguous classification of the chimpanzee. Less than 20 years later, in 1775
Johann Friedrich Blumenbach gave the species its final scientific name: Pan troglodytes.
11
Another “type” of chimpanzee was scientifically discovered in the 1910s-1920s (Coolidge &
Shea, 1982). This different “variety of chimpanzees”, as first described, was separated from
Pan troglodytes in 1933 by Harold Coolidge Junior, and named Pan paniscus (Coolidge,
1933). This species is physically different from the other, being qualified as more gracile and
relatively shorter than Pan troglodytes (Coolidge, 1933; Coolidge & Shea, 1982). Therefore,
soon after its discovery Pan paniscus was called “pygmy/gracile chimpanzee”, to differentiate
from Pan troglodytes which was called “robust chimpanzee”. For the sake of clarity, I will
use the common names chimpanzee and bonobo respectively for Pan troglodytes and Pan
paniscus.
Chimpanzees and bonobos exclusively occur in Africa. They are considered to be the closest
living relatives to humans. An average of 98.8% of genes is shared between Homo spp. and
Pan spp. (Furuichi & Thompson, 2008). The last common ancestor of these two genera was
estimated between 5 and 7 MYA (Dawkins, 2004). The estimation of the split between
chimpanzees and bonobos locates their last common ancestor much more recently: between
0.68 and 2.1 Ma (0.68-1.54 Ma: Hey, 2010; 0.86-0.89 Ma: Won & Hey, 2005; 1.5-2.1 Ma:
Stone et al., 2010). After the divergence of these two species of Pan, the bonobo was
restricted to the rainforest located between the Congo and Kasai rivers in the Democratic
Republic of Congo (Dawkins, 2004), while the chimpanzee conquered the forested regions of
the lowest latitudes of the African continent (Hey, 2010).
2.2. Pan troglodytes
2.2.1. How many subspecies?
Pan troglodytes is the most widespread of all great apes, and probably the haplorhine primate
with the largest distribution area in the African tropical forests (Oates, 2006) (the tantalus
monkey (Chlorocebus tanalus) known as the primate with the largest range (Rowe, 1996) is
little represented in forests). The species occurs in more than 18 African countries, from
Senegal to Tanzania (Fowler & Sommer, 2007). Throughout their wide distribution crossing
several taxonomic barriers, chimpanzees have been divided into several subspecies based on
geographically, genetically and physically differences (Mwambo, 2011). Nevertheless, the
number of subspecies, as well as their characteristics, has been difficult to determine, and the
taxonomy is controversial.
12
For 80 years, three subspecies of chimpanzees were accepted (Morgan et al., 2011), mostly
based on their geographical distinctiveness. As its name suggests, the Western chimpanzee
(Pan troglodytes verus) is the subspecies with the most Western geographic range. This
chimpanzee occurs from Senegal to Ghana but is locally extinct in Gambia and possibly
extinct in Benin, Burkina Faso and Togo (Oates et al., 2008). Some populations of this
subspecies undergo the driest conditions of the natural home range of Pan troglodytes. While
most chimpanzees tend to live in humid areas, the well-studied population of Pan troglodytes
verus of Fongoli (Senegal) subsists in a semi-arid mosaic savannah habitat with a small
portion of gallery forest (only 2% of their home range) and an average rainfall of 800-
1000mm/year (Pruetz & Bertolani, 2007). The central chimpanzee subspecies has been
recorded between Cameroon and the Ubangui River/Congo River in Congo (Mwambo, 2010).
It is the most common subspecies (Stone et al., 2010). The third subspecies presents the most
Eastern distribution, being located between the East of the Ubangui River/Congo River and
Western Tanzania: the Eastern chimpanzee (Pan troglodytes schweinfurthii) (IUCN, 2013).
Small relict populations of this taxon are still remaining in Burundi and South-Eastern Sudan
(Oates et al., 2008).
The genetic differences between the subspecies of Pan troglodytes have led to singular
phenotypes: Eastern chimpanzees tend to have longer hair and bronze to copper facial skin
(Mwambo, 2010). On the other hand, central chimpanzees have a dark pink face and lose the
hair of their forehead as they grow old, and the dark area around the eyes of the Western
chimpanzees is the reason why this taxon is sometimes called “the ape with glasses”
(Mwambo, 2010).
The taxonomic status of the population of chimpanzees restricted to the South and the East of
Nigeria and Western Cameroon (until the Sanaga River) is subject to controversy. A growing
international interest for this chimpanzee started quite recently, after the publication of the
research of Gonder et al. (1997). In this study, not only was the Nigeria-Cameroon
chimpanzee recognized as a distinct subspecies, but also the most genetically differentiated
taxon, based on mitochondrial evidence. The dental morphometric research of Pilbrow
(2006), and the genetic research of Ghobrial et al. (2010) both supported the differentiation of
this population of chimpanzees in a distinct subspecies. This chimpanzee was first called Pan
troglodytes vellerosus from the collection records of a specimen collected in 1862 by Richard
Burton from Mt. Cameroon (Hey, 2010; Oates et al., 2009). Further studies on the archives of
this individual have shown this specimen was actually from Gabon (South-East of the Sanaga
13
River), and therefore actually belonged to the central chimpanzee taxon (Oates et al., 2009).
Hence, the name of this subspecies was then changed into Pan troglodytes ellioti (Matschie,
1914). But the distinction of this taxon is still controversial and subjects to disagreements
(Fisher et al.¸ 2006; Morgan et al., 2011). Some specialists, using Y-chromosome evidence,
argue the populations of chimpanzees of Nigeria and western Cameroon are not differentiated
enough to be recognized as a singular subspecies (Stone et al., 2002) while others argue Pan
troglodytes ellioti is genetically the most distinct population of chimpanzees (Gonder et al.,
2011). The absence of significant differences between Nigeria-Cameroon chimpanzees and
Western, central and Eastern chimpanzees might however come from a lack of sufficient data
on individuals from the Bight of Biafra (Stone et al., 2010; Taylor & Groves, 2003). More
research is needed to define the affiliation of the isolated population of chimpanzee of South-
Western Nigeria to either Pan troglodytes ellioti or Pan troglodytes verus (Gonder et al.,
2006; Greengrass, 2009; Morgan et al., 2011).
The controversy in Pan troglodytes taxonomy is not only restricted to the status of the
Nigeria-Cameroon chimpanzee, and is largely dependent on the phenotype or genetic
sequencing on which is based the comparison. Pan troglodytes marunguensis has been
proposed as a 5th
subspecies to detach the population of chimpanzees of the West of
Tanganyika Lake from Eastern chimpanzees (Grubb et al., 2003). More recently, Gonder et
al. (2011) showed the evidence of genetic differences which would separate the chimpanzees
from Southern Cameroon into a distinct population. Morin et al. (1994) have proposed the
separation of Pan troglodytes verus from the other subspecies into a completely distinct
species based on mitochondrial DNA sequencing. In my thesis, I will use the classification of
chimpanzees proposed by the systematics of African primates published by Grubb et al. in
2003: four subspecies known as the central (Pan troglodytes troglodytes), Eastern (Pan
troglodytes schweinfurthii), Western (Pan troglodytes verus), and Nigeria-Cameroon
chimpanzees (Pan troglodytes ellioti). The complete distribution of the two species of Pan sp.
(chimpanzee and bonobo) and the four subspecies of Pan troglodytes spp. is shown in Figure
2.
14
LEGEND :
1. Distribution of Pan sp. :
: Pan paniscus
: Pan troglodytes ellioti
: Pan troglodytes schweinfurthii
: Pan troglodytes troglodytes
: Pan troglodytes verus
: Pan troglodytes spp. – isolated metapopulation needing further research to be classified either P. t. verus or
j P. t. ellioti.
?: Countries where chimpanzees have possibly gone extinct.
2. Sympatric zones:
1: Sanaga River – proposed taxonomic border between P. t. t. and P. t. e..
2: Mbam River
3: Niger River – potential barrier between the East-Nigerian and the South-western Nigerian P. t. spp. populations.
4: Congo/Ubangui River – taxonomic border between P. t. t., P. t. s. and P. p.
: Gene flows between P. t. t. and P. t. e. - Hybridization area.
: Gene flow between P. t. t. and P. t. s. - Hybridization area.
3. Study Site:
? ?
?
1000 km
600 miles S
N
W E
100 km
75 miles
4
1
3
2
1
Figure 2: Distribution of Chimpanzees and Bonobos based on Butynski, 2001; Gonder et al.,
2011; IUCN, 2013; Rayner et al., 2011.
15
2.2.2. Behaviour and Ecology
Chimpanzees are characterized by a highly complex social life and variation between groups,
in tradition, technology, feeding, and behaviour which is described by a growing scientific
group as cultural differences (Fowler & Sommer, 2007; Reynolds, 2005). Across the species
distribution, the use of tools has been recorded in more than 25 different contexts (Pruetz &
Betolani, 2007). Some technologies are restricted to populations, or even groups (such as the
creation of spears to hunt galagos (Galago spp) which has only been recorded in the
population of Fongoli: Pruetz & Bertolani, 2007), while other are widespread throughout the
home range of chimpanzees. However, even if they use the same technology, different groups
of chimpanzees may use different techniques. For example, although the technology of insect-
fishing has been recorded in Nigeria-Cameroon (Fowler & Sommer, 2007), Eastern
(Reynolds, 2005), central and Western chimpanzees (Humle & Matsuzawa, 2002) its
technique of application varies. Between groups of chimpanzees, many differences have been
recorded including the handling-style, the size and the number of sticks, as well as the
position of the “fisher” (on a branch or on the floor), and the targeted insects (Humle, 2013;
Schöning et al., 2008).
The spreading of such tradition and culture is still unknown. The nut-cracking behaviour
largely reported in Pan troglodytes verus populations since 1844, had never been observed in
any other subspecies until 2006 (Wrangham, 2006). Morgan & Abwe (2006) recorded this
behaviour in a Pan troglodytes ellioti population of the Ebo Proposed Protected Area (South-
West Cameroon), more than 1700km away from the most Eastern smashing-nut site (Ivory
Coast). In contrast, the nut-cracking behaviour has never been reported since the beginning of
the long-term study of Nigeria-Cameroon chimpanzees in Gashaka-Gumti National Park,
Nigeria (Fowler et al., 2011). Other activities such as nest building have presented variations
between populations related to both available vegetation, and culture (Fruth & Hohmann,
1994).
Given the rate of the destruction of the natural habitat of Pan troglodytes and the heavy
hunting pressure the species undergoes throughout tropical Africa, it is likely that the
diversity of chimpanzee culture will decrease rapidly before its richness is scientifically
recorded (Reynolds, 2005).
16
2.2.3. Conservation Status
The four subspecies of chimpanzees have been red listed as endangered taxa by the IUCN
since 1996, and live in more and more fragmented populations (Balcomb et al., 2000). Junker
et al. (2012), suggest that the suitable environmental conditions for Pan troglodytes verus
were reduced by 11% between the 1990s and the 2000s. The same study showed the central
chimpanzee subspecies lost 17% of its suitable habitat within the same two decades.
Chimpanzee populations are facing dramatic declines, including in countries where they have
been historically recorded as living in large populations, such as Ivory Coast and Gabon
(Greengrass, 2009). Alarmed by the fast decreasing of the wild populations of chimpanzees in
Congo and Gabon (the two countries believed to possess the largest populations of great
apes), Walsh et al. (2003) proposed to reinforce the protection status of the species and to red
list it as „Critically Endangered‟. This statement, however, has been criticized by many
specialists. Oates (2006) argued chimpanzees were less endangered than gorillas and orang-
utans (Pongo spp.). He stated, even if by strict application of IUCN threat criteria, Pan
troglodytes can be considered as endangered (slow development of infants and significant
hunting pressure), it is unlikely to go extinct by 2100. Nevertheless, this statement does not
mean chimpanzee populations are not under unsustainable human pressures.
With a population of chimpanzees composed of more than 30,000 Pan troglodytes troglodytes
and more than 3,000 Pan troglodytes ellioti (Ngalla e al., 2005) sharing sympatric zones
where gene flows enrich each taxon (Gonder et al., 2011), Cameroon is one of the most
important countries in terms of chimpanzee diversity. Conscious about this richness, the
Government of the country elevated both subspecies into A class animals, prohibiting the
hunting, trade and use as pet of chimpanzees (Djeukam, 2012). However, in the last decades,
there have been an increasing number of resident chimpanzees in Cameroonian wildlife
rescue and rehabilitation centres such as Limbe Wildlife Center, Mefou National Park, and
Sanaga-Yong Chimpanzee Rescue Center (Ghobrial et al., 2010). These orphans, victims of
illegal hunting, represent an indirect measurement of the extent and the unsustainability of
poaching throughout the country. Ghobrial et al. (2010) used genetics to establish the map of
the origin of chimpanzees rescued in Limbe Wildlife Centre. This map showed Pan
troglodytes ellioti and Pan troglodytes troglodytes are hunted throughout their whole range in
Cameroon. More chimpanzees coming from the North of the Sanaga River (including some
places in Nigeria) than the South of the river were found, certainly because the rescue centre
is located in Limbe which is in the South-West region of Cameroon. Unlike well-organized
17
elephant tusk networks which operate internationally, the traffic of dead and alive
chimpanzees is therefore described as local and widespread.
Chimpanzees suffer from their title of „close relative to humans” as well as their local
traditional image. While the gorilla represents strength and power (used as an antibiotic,
gorilla bones are believed to make children stronger in South-Western Cameroon: Etiendem
et al., 2011) in many Cameroonian tales, the chimpanzee is seen as an intelligent, clever
human-like primate (Mfomo, 1982). This picture of human-like primate puts the chimpanzee
as one of the “finest” animals for traditional medicine. For example, in 1994, King stated
some tribes in the South-West region of Cameroon heal sprains and breaks by applying boiled
chimpanzee bones.
According to White & Edwards (2000), gorillas are still found in normal population sizes in
secondary forests, but in contrast, chimpanzees strongly prefer primary forests and only
recolonize disturbed areas long after the logging event. However, a study made by Matthews
& Matthews and published in 2004 showed chimpanzees are not always displaced by logging.
The results of this study conducted in both protected and logged areas in and around the
Campo and Ma‟an forests in Southern Cameroon, showed as long as the habitat is a mosaic of
primary and secondary forest, and the other anthropomorphic activities (especially hunting)
are prevented, chimpanzee density does not automatically decline.
The consequences of human activities upon chimpanzee populations are especially alarming
for the Nigeria-Cameroon chimpanzees. Farming (the main threat in the Bamenda Highlands:
McKay & Coulthard, 2000), hunting, cattle grazing, trading to zoos and American
laboratories are happening too often and in too many areas across the range of Pan
troglodytes ellioti to be sustainable for the populations (Mwambo, 2010).
2.3. Pan troglodytes ellioti
2.3.1. The Most Threatened Subspecies of Chimpanzees
The Sanaga River has been proposed as the natural barrier between the Nigeria-Cameroon
chimpanzee and the central chimpanzee (Fowler & Sommer, 2007; Morgan et al., 2011)
(Figure 2). This border is however not strict as some limited gene flow has been reported
between these two subspecies of chimpanzee in the Centre region of Cameroon, around the
confluence of the Sanaga and Mbam rivers (Ghobrial et al., 2010) (Figure 2).
18
With a total of 3,500-9,000 individuals in the wild and only 70 in Cameroonian and Nigerian
sanctuaries (Morgan et al., 2011), the Nigeria-Cameroon chimpanzee is the most threatened
subspecies (Fowler et al., 2007; Ghobrial et al., 2010), having both the least population size
and geographic distribution (Mwambo, 2010). The largest numbers of chimpanzees are found
in the South-West region of Cameroon (Morgan et al., 2011). The only healthy populations of
Pan troglodyte ellioti in Nigeria are found in Gashaka-Gumti National Park which possesses
several hundred individuals (Fowler et al., 2007). In Cameroon, the most important
populations are located in the proposed Ebo National Park (probably more than 750), in
Mbam and Djerem National Park (at least 500) and in Banyang Mbo sanctuary (500-1,000)
(Morgan et al., 2011).
Species that are rare either in terms of their distribution or population densities are generally
more vulnerable to extinction risks compared to species that are not rare (Gaston, 1994). Pan
troglodytes ellioti falls into this category, therefore an improvement of the local conservation
of this subspecies is urgently needed. The populations of this endangered great ape are
becoming more and more fragmented into isolated sub-populations throughout most of their
former range (Mwambo, 2010). Using the population viability analysis tool VORTEX, the
most pessimistic estimations of Hughes et al. (2011) estimates the extinction of Pan
troglodytes ellioti before 2035. In order to prevent this expected loss of diversity in the Pan
genus, the IUCN recently published the “Regional Action Plan for the Conservation of
Nigeria-Cameroon Chimpanzee (Pan troglodytes ellioti)” (Morgan et al., 2011). In this action
plan were proposed research and managements to improve the protection of the subspecies in
both Cameroon in Nigeria. Because the estimation of the size of the population of a taxon is
essential for good conservation practice (Beck & Chapman, 2008), this action plan
recommends several priority survey sites: four in Nigeria and six in Cameroon. In these six
Cameroonian proposed survey sites, two are situated in the North-West region, one being the
forest of Tubah Sub-Division. The presence of chimpanzees in this forest has long been
verified (Ngalla et al., 2005), but no scientific research has been conducted to estimate the
population of great apes.
2.3.2. “The Great Apes of Tubah”
Little is known about the chimpanzees of Tubah Sub-Division. For about ten years, an
informal chimpanzee monitoring has taken place. This work has been sponsored by John
19
DeMarco, a Canadian citizen, who previously worked on the „Kilum-Ijim Forest Project‟, and
the “Bamenda Highlands Forest Project”, both dedicated to the sustainable management of the
montane forests. When these two projects ended in 2003, he decided to start an informal
monitoring program on the chimpanzees of the Bamenda Highlands. This monitoring has
been done regularly starting first with Romanus Ikfuingei (now working for the Wildlife
Conservation Society in Limbé, South-West Cameroon), and then with two part-time
monitors supervised by Ernest Vunan. Unfortunately, the lack of additional support and
scientific knowledge has prevented efficient monitoring: the monitoring is random, the
animals are not followed-up, and all the patches are not supervised. Nonetheless, these
activities have certainly had a positive impact on the conservation of the chimpanzees of the
forest, showing the villagers about the necessity of protecting the apes. Moreover, because of
the observation of at least one baby, it was established Pan troglodytes ellioti were living in a
breeding group. Additional attention was given to the chimpanzees of Tubah in 2008-2009,
during a local awareness campaign focusing on the protection of “The Great Apes of Tubah”
by an NGO based in Bambui: the Benevolent Association for the Protection of the
Environment and Socially Underprivileged (BAPESU). Unfortunately, this campaign was
planned to exclusively inform villagers of Bambui despite the fact the fragment of forest
belonging to this village isn‟t visited by chimpanzees anymore due to an important
anthropomorphic presence. In 2011, a local villager was condemned to serve a couple of years
in prison for the killing of a chimpanzee. As a result of this success in the fight against
poaching, villagers and hunters are now reluctant to kill chimpanzees.
A recent study conducted by Bergl et al. (2012) expanded the known distribution of the
Critically Endangered Cross River gorilla and highlighted the importance of corridors to
prevent populations from isolation. The forest of the subdivision of Tubah could be an
essential corridor between bigger undisturbed areas with healthier chimpanzee populations
such as Kom-Wum Forest Reserve.
3. Estimating the Population of Chimpanzees
According to Beck & Chapman (2008), it is essential to know the average population of an
animal to protect it. Many methods have been created since the scientific concern of counting
wild animals. This practice presents some problems in forested areas, where taxa are more
20
difficult to find than in open environment such as savannahs. Because of their natural habitat
mostly composed of primary or secondary forests, the number of individuals in a group of
chimpanzees is more often estimated than certain. Moreover, chimpanzees occur in relatively
low density, which makes it difficult to observe them directly (Matthews & Matthews, 2004).
In his book about the population of Pan troglodytes schweinfurthii of the Budongo forest
(Uganda) he studied for decades, Reynolds (2005) stated it took his research team several
years to accurately count one group of chimpanzees using the habituation method. The
habituation method is the most efficient method to know the overall population of great apes,
but it is particularly difficult, expensive and time consuming.
The nesting behaviour of chimpanzees is extensively discussed in the literature (Beck &
Chapman, 2008; Blom et al., 2001; Last & Muh, 2013). Because of the slow decay rate of a
nest (highly variable: from 73 to 221 days: Kühl et al., 2008) and therefore the long-lasting
presence of nest sites, the counting of night nests of chimpanzees is the most popular
technique (Balcomb et al., 2000; Kühl et al., 2008; Matthews & Matthews, 2004; White &
Edwards, 2000).
Several types of methodology have been used for chimpanzee nest counting. The two
methods the most used to conduct such surveys have been the „transect method‟ and the
„recce walk method‟, also called „reconnaissance walk method‟ (Kühl et al., 2008; Ross &
Reeve, 2010; White & Edwards, 2000). The latter is mostly used during pre-studies to have a
first glance of the distribution of the apes.
While surveys on populations of chimpanzees are numerous in the literature (Reynolds, 2005;
Walsh et al., 2003; Yoshikawa et al., 2008), most of them have been restricted to the central,
Western and Eastern subspecies with only few studies made on Pan troglodytes ellioti
(Ghobrial et al., 2010). Most of the studies on the Nigeria-Cameroon chimpanzee have been
conducted since the distinction of the taxon as a separate subspecies (within the last two
decades). Out of these few surveys and research on the least known chimpanzee, most of the
work has been done in Gashaka-Gumti National Park in Nigeria, by the “Gashaka Primate
Project” (Beck & Chapman, 2008; Fowler et al., 2007; Sommer et al., 2004) and in the Ebo
Proposed Protected Area (Abwe & Morgan, 2008; Morgan & Abwe, 2006).
Although the North-West region of Cameroon is in the middle of the range of the Nigeria-
Cameroon chimpanzee, no scientific survey has been conducted on this endangered primate
and little is known about its distribution in the region.
21
4. The Study
4.1. The Montane forests of Tubah Sub-Division
The landscape of the North-West region of Cameroon has been shaped by the Cameroon line.
About 75% of the province is situated above 1000m (Fongjong, 2008). The Bamenda
Highlands are composed of peaks up to 3008m (Mt Oku), a high lava plateau between 2000
and 2800m, an intermediate plateau between 1000 and 1900m, and a low plateau between 300
and 900m of altitude (Ndenecho, 2009). These mountains are made up of volcanic products
(alkali basalts, phonolites, trachytes and ignimbrites) and age from 17.4 Ma to present
(Gountie Dedzo et al., 2011).
One of the remaining forest blocks of these mountains is the forest of Tubah Sub-Division,
surrounding the villages of Bambui, Babanki (divided in two villages: Kedjom-Keku and
Kedjom-Ketingo) and Finge (Figure 3). Located 17km from Bamenda (the largest city of the
region), this forest does not have a common name (the name “Tubah Forest” is attributed to a
Kedjom-Keku
Bambui
Bambili
Sabga
Finge
S
N
W E
Douala
Yaoundé
Bamenda
1000 km
600 miles
Kedjom-
Ketingo
Northern Borders of
Bamenda
2.5 km
1.5 miles
Figure 3: Satellite map of the Forest surrounding the villages of Kedjom-Keku and
Kedjom-Ketingo (Babanki), Bambui and Finge (green polygon). Although placed on
the map, Bambili and Sabga do not possess any patch of the forest covered in the
study site. Picture taken from the software Google Earth©.
22
non-native forest of Eucalyptus sp.). On the advice of John DeMarco, the name Babanki-
Finge was chosen, using the two villages most concerned.
4.2. Aims & Objectives
This study has three aims:
1) To estimate the population of Nigeria-Cameroon chimpanzees and record the species
of monkeys in the Babanki-Finge Forest
This is the first study of the haplorhine primate abundance and distribution in the Babanki-
Finge Forest. The presence of the rare Nigeria-Cameroon chimpanzee and several species of
monkeys have been witnessed by local villagers (Ingram & Nsom Jam, 2007), but a scientific
survey on the population of Pan troglodytes ellioti is needed (Morgan et al., 2011). Moreover,
the presence of a population of Preuss‟s guenons would dramatically increase the interest of
Babanki-Finge.
2) To map the Vegetation of the Babanki-Finge Forest
The only available map of the Babanki-Finge area is the map of the villages surrounding the
forest, dating of 1913, and the satellite map. It is therefore essential to create the map of the
forest for a better knowledge of the region.
3) To record the Human Pressure Upon the Babanki-Finge Forest
The forest of Babanki-Finge is an unprotected forest fragmented in several patches. Due to its
lack of protection status and its location within the limits of four expanding villages, the forest
is subject to anthropomorphic pressure. Evaluation of this may help determine the relative
importance of different threats to this forest.
23
Chapter 2 – METHODS
Detailed maps have widely been regarded as essential for a survey (Buckland et al., 1993).
However, no maps of the forest of Babanki-Finge have ever been created. Therefore, the first
purpose of this study was to create a map of the forest, detailed enough to differentiate the
various patches of woodland from the land used for cattle-grazing and farms. As a
consequence, on the field, the data collection was made in two steps: first the delimitation of
the forest, and then the recording of signs of haplorhine primates.
1. Study Site
Babanki-Finge is a fragmented montane forest divided in sets of patches (Figure 4).
Therefore, the limits of the study site were difficult to define. It was decided to use the
borders delimiting the area visited during the informal monitoring of chimpanzees. Babanki-
Finge is located within the borders of three villages: Bambui, Finge, and Kedjom-Keku
(Figure 4). The latter is also called Big Babanki, to differentiate with Small Babanki, also
called Kedjom-Ketingo. These two villages share a common and relatively recent history.
According to the oral history of the villages, Babanki used to be one single village located in
the current site of Small Babanki. Less than 100 years ago, the villagers of Babanki separated
and built two villages: Kedjom-Keku („People of the forest‟) and Kedjom-Ketingo („People of
the mountain‟). Therefore, the village borders inside the forest are not well determined.
Nonetheless, because of the closeness and relatedness between the two villages, and the
uncertainty of the borders between them, I decided to include both Small and Big Babanki in
my thesis and to simply refer to them as Babanki.
Babanki-Finge is a small forest fragmented by grassland patches. The climate provides four
seasons: one long and one short dry season (December, January, and February are the driest
months), and one long and one short rainy seasons (July and August are the wettest months)
(Ndenecho, 2011). The mean temperature is roughly comprised between 13°C and 22°C, with
an average of 1,780-2,290mm/year of rainfall (Ndenecho, 2011).
24
2. Data collection
2.1. Study Period
I worked in the Babanki-Finge forest from the 25th
of February to the 15th
of May 2013. This
11.5-weeks field work started with a 3-day pilot study. During this short period, two methods
of data collection were used and compared: the “transect method”, and the “reconnaissance
Kedjom-Keku
Finge
Bambui
Kedjom-Ketingo
ALEGAFOR
Kubuh
Mendong I
Fintse
Mendong
II
KEFFEM
ABONG-PHEN
2.5 km
1.5 miles
S
N
W E
Figure 4: Satellite map of the Babanki-Finge forest. The three ensembles of patches
covered in this study are encircled in green. The sub-patches are encircled in yellow.
The dashed line divides the two satellite pictures: the most Northern one was taken the
31st of January 2010, and the most Souhern one was taken the 13st of June 2011.
Picture taken from the software Google Earth©
25
walk method” (or “recce walk”). Despite its wide use (Beck & Chapman, 2008), the distance
transect method presents disadvantages which turned to be exacerbated on this study site due
to the characteristics of Babanki-Finge. First, although several factors essential for the
transect method (such as the length of line transects, their orientation, etc.) should be based on
data from pilot studies (Buckland et al., 1993), no studies related to the haplorhines, or any
other mammals have been done in this forest. In his guide for transect methodology, Peres
(1999) suggested that 4-5km transects for a data collection should be sufficiently
representative. He also stated transects in areas smaller than 500ha (i.e. areas such as the
Babanki-Finge forest), should be cut parallel to each other. These conditions were not feasible
in my study site because the sizes of the forest patches were far too small to cut transect lines
longer than 2km, and even if it were possible, the rugged landscape prevents any parallel lines
to be formed. Moreover, the creation of straight lines cut in the heart of the forest requires a
small (but not negligible) destruction of the habitat (White & Edwards, 2000), which must be
proscribed in Babanki-Finge, due to the large fragmentation of the forest and the high degree
of hunting. Therefore, the “recce walk” method was preferred for this study (see below for
details of methods).
An important part of the data collection was done in collaboration with the local team
monitoring chimpanzees. On the 10th
of April, after a meeting with John DeMarco (sponsor of
the monitors), a schedule was established to work in the field with David Achomochi and
Moses „Pa‟‟ Amah Alukeh, and their supervisor Ernest Vunan. Their skills and knowledge of
the forest became indispensable for moving through the dense vegetation, discovering signs of
chimpanzees, and naming tree species. During the surveys occurring in two of the most
remote patches (Mendong I and Abong-Phen), we were lodged in one of the two stations used
by the monitors. In one of our weeks in the field, Julius Mbicho Abongeh joined our team as a
photographer.
After my departure from the study site, Ernest and the monitors decided to complete the data
collection by doing recce walks in the last two unexplored remaining patches: Kubuh and
Mendong II. Lacking measurement devices, they only recorded the age of the nests, the tree
species and the GPS location for each nest. This extends my data collection to the 6th
of June.
26
2.2. The Global Positioning System (GPS)
Initially created for military applications, the GPS is a satellite-based navigation system
composed of 24 satellites placed into orbit by the United States Department of Defense
(DOD) and operated by the United States Air Force 50th
Space Wing (Ta, 2011). According to
LeJeune & Rondeux (2007), the Garmin GPSMap 60CSx is a very reliable device, with a
10m-precision under the canopy of the forest. Therefore, this model was chosen for this study.
The 2-dimensional Cartesian coordinate system used was the Universal Transverse Mercator
(UTM). The device was set on 32N UTM, which corresponds to the zone where the North-
West region of Cameroon is located.
The data collection was mainly done using two features of the device: “Tracklog” and
“Waypoints”. The first element was programmed to record the location every 50m, creating a
track later transferred to the software Garmin® MapSource®. It allowed recording completed
roads during the delimitation of the forest. It was also used during the recce walks in order not
to use the same path twice, or to accurately map some distinct areas within the forest (e.g.
farms, open area with Aframomum sp., etc.). Unlike the “Tracklog” feature, the “Waypoints”
program allows the researcher to mark independent points. In my study, it was used during
forest patch delimitations and recce walks to both record any changes in the environment and
to note the location of discovered chimpanzee nests.
2.3. Forest Delimitation
The GPS device was used to record the track delimitating the edge of the forest. If a small
farm or some dense vegetation prevented us from moving directly on the edge of the
woodland, the distance between the device and the forest was recorded. When a rugged land
or precipitous slopes did not permit us to walk either outside or inside the border of the forest,
photographs or videos were taken, and pictures were drawn to capture characteristics of the
shape of the forest.
27
2.4. Diurnal Primate Signs
Monkeys and apes were recorded via direct and indirect signs. Direct signs were not expected
to be frequent due to the close proximity to villages and the high degree of anthropomorphic
pressure (mainly farming and cattle grazing). Indirect signs specifically targeted ape nests.
2.4.1. Direct Signs
Direct observations of monkeys and apes were recorded, as well as vocalisations. For each
observation, the location, the time, the species, the number of seen individuals, and the
number of estimated individuals were noted.
2.4.2. Indirect Signs
Indirect signs of haplorhine primates were also recorded and the location of each sign was
noted.
Indirect Signs - Nest Recording
Gorillas and chimpanzees both build ground and tree nests which can be confounded (White
& Edwards, 2000). Nevertheless, the current distribution area of the Cross River gorillas (the
only subspecies of gorilla located on the Nigeria-Cameroon border: Bergl & Vigilant, 2007)
does not include the Bamenda Highlands. In the beginning of my field work, a silverback
gorilla was shot dead around Santa (a few kilometres from Bamenda), but the presence of
Gorilla gorilla diehli that North is exceptional and restricted to isolated individuals.
Therefore, all discovered nests were assumed to be built by chimpanzees.
For each nest, we recorded the geographic position (using GPS), the height (when
measurable), the tree species and the estimated age of the nest. The tree species were only
recorded in presence of the field assistant Ernest Vunan who acted as the botanist of the team
due to his great experience in the Babanki-Finge forest and his knowledge in plants. The
height was measured as shown in Figure 5. The age of the nests was estimated using Table 1.
28
Figure 5: Drawing representing the requisite measurements for the assessment of the
height of a chimpanzee nest. I was the only observer, therefore the distance between my
eyes and the ground is 1.87-0.10=1.77m. The trigonometric formula was used to measure
the height of the nest in the tree.
Table 1: Descriptive of the estimation of the age of nests.
Age
Observations Estimation
green leaves, thick nest Fresh
green and brown leaves Recent
brown leaves Old
few brown leaves and visible branches - can see
through the nest Very Old
B
C
A
𝑡𝑎𝑛 𝐵Â𝐶 =𝐵𝐶
𝐴𝐵 <=> 𝐵𝐶 = 𝐴𝐵 × tan𝐵Â𝐶
1,77m
Distance
Tree – Collector
Angle
29
Indirect Signs – Other Signs
The lack of knowledge about the ecology of the chimpanzees and monkeys of the Bamenda
Highlands (such as the species of trees on which they feed on, their technology used, etc.)
limited us in the collection of signs other than nests. Therefore, we did not focus on searching
for other signs (e.g. faeces, tracks, footprints, rests of fruits), even though they were recorded
when observed.
2.5. Habitat Description
The description of the habitat was done progressively, during both the delimitation of the
limits of the forest, and the recce walks. The vegetation types were categorized as shown in
Table 2. Human settlements are very old in the region around Bamenda, as well as the
anthropomorphic disturbance, and the Babanki-Finge forest has supported much human
pressure throughout the last centuries. This constant disturbance can be noticed in some
patches where aligned knolls (characteristics of farming) are found with mature indigenous
trees, in the middle of the forest, recovered by the vegetation of the understorey. Idecided not
to use the terms “primary” and “secondary” forests for my study, partly because the definition
of primary and secondary forests is based on a potentially ambiguous estimation of the
maturity of trees (Chokkalingam & De Jong, 2001), and also because it was not possible to
estimate the age of the recovered forest. Instead, a sub-categorization of the forest was used
(Table 3), based on the vegetation descriptive presented in the study of De Vere et al. (2011).
Figures 6 and 7 are photographs of typical plants of the riparian forest: the spiny tree fern
(Cyathea manniana), and an unknown species of tree.
The canopy cover was assessed using the descriptive used by Wiseman (2011) in work in the
Kagwene Gorilla Sanctuary, Cameroon (Table 3). The estimation was done by forming a
circle with the thumb and the index of both hands. Placed under the canopy, the observer
looked up and evaluated the percentage of cover. I was the only one to do the observations
and to record the canopy cover estimation, in order to prevent any observer variation in the
data collection. The angle of the slope of the forest was also estimated, using Table 3.
Finally, the average height of the canopy was recorded and classed in to categories: below or
equal to 10m, and above to 10m. This height was chosen in order to better differentiate two
classes of forests: young and mature forests. Mature forests of the Bamenda Highlands are
30
composed of tall trees such as Carapa grandiflora (Meliaceae) and Prunus africana
(Rosaceae) of which adult individuals can reach 30m and above (Farwig et al., 2006; Ingram
& Nsom Jam, 2007).
Table 2: Vegetation Descriptive based on De Vere et al., 2011.
Vegetation Type - Code Description
Forest – FR Vegetation dominated by young to mature trees, with an
understorey more or less dense – Carapa grandiflora,
Prunus africana, Strombosia sp., Ficus sp., etc.
Riparian Forest - RF Type of forest directly bordering streams, dominated by
Cyathea manniana (Figure 6), and a short unknown tree
covered with moss (Figure 7).
Secondary Vegetation – SV Vegetation dominated by opportunistic/fast-growing
plants, no mature trees.
Aframomum – AF Vegetation dominated by Aframomum sp.
Gnidia glauca – GG Clear forest dominated by Gnidia glaucau.
Acacia sp. – AC Clear forest dominated by Acacia sp.
Shrubs – SH Mix of bush and tall grass - Erica manni, Eugenia gilgii,
etc.
Fern – FN Habitat dominated by Pteridium aquilinum spp.
Tall Grass – TG Habitat dominated by tall grasses – Lobelia columnaris,
Pennisetum purpureum, etc.
Grassland – GL Grazing land characterized by short-grass and cattle
paths.
Farmland
Farm – FM Mono- or poly-cultures of crops – Zea mays, Dioscorea
rotundata, Solanum scabrum, Rhapia mambillensis, etc.
Disturbed Forest – DF Forest with understorey composed of farms – Colocasia
esculenta, Coffea sp., etc.
Ancient Farm - AF
Open area within the forest, characterized by a growing
vegetation slowly replacing mono- or poly-cultures –
Persea ammericana, Musa sp., etc.
Eucalyptus – EU Mono-culture of Eucalyptus sp.
31
Table 2 (following): Vegetation Descriptive based on De Vere et al., 2011.
Vegetation Type - Code Description
Burnt Area – BA Open area recently cleared for the creation of
a new farm.
Rock – RC Large rock within or outside the forest,
without any vegetation on top.
Stream - ST Small to large river, at least 50cm wide (the
distance 50cm was chosen because the data
collection started at the end of the dry
season, when the width of streams is
significantly shortened)
Table 3: Sub-categorization of the forest descriptive.
Sub-categories Characteristics
Canopy cover
0 0-25% coverage: no canopy/open canopy
1 26-50% coverage
2 51-75% coverage
3 76-100% coverage: closed canopy
Slope
0 Flat Ground
1 0-10% (0-5.7°): gentle slope
2 10-50% (5.7°-26.6°): steep slope
3 >50% (>26.6°): precipitous slope
32
Figure 6: Photograph of a mature Cyathea manniana.
Figure 7: Photograph of an unknown species of trees, typical of the Riparian Forest in
Babanki-Finge.
33
2.6. Fig Trees
The importance of fig trees (Ficus sp.) in the diet of chimpanzees and other tropical primates
has largely been documented (Tweheyo & Obua, 2001; Wrangham et al., 1991). Therefore, I
originally intended to record the number of fig trees in each forest patch. However, in mid-
April, when I started working with Ernest Vunan, I learnt there are several species of fig trees
in Babanki-Finge, and not all of them are used by the chimpanzees and monkeys. As I did not
have the expertise, nor time, to distinguish the different species of trees, I did not
systematically record fig trees.
3. Data Analyses
3.1. Creating a Map
The recovery, storage, and display of geographic information were completed using
Geographic Information Systems (GIS) (Rogerson & Fotheringham, 1994). The software used
for analysing and exploiting the collected data was ArcGIS version 9.1. In order to plot the
collected points and to map the vegetation accurately, satellite pictures from Google Earth©
were georeferenced into ArcGIS. Two satellite pictures of the Babanki-Finge forest were
taken from Google Earth©. The picture covering most of the forest was taken on the 31st of
January 2010, while the picture representing the extreme South of the forest (the patch
belonging to Bambui named Alegafor) was more recent, dated 13/06/2011. Despite their date,
the photographs matched the collected data (Figure 8), with only small irregularities.
In addition to waypoints and track logs, pictures, videos and drawings were used to complete
the map in areas where precipitous slopes did not allow data collection.
Although no mapping was done in Kubuh and Mendong II (the two patches covered by the
field assistants when I had already left Cameroon), I decided to add these two forested areas
in the map of Babanki-Finge. Because of the apparent few differences between the satellite
images and the GPS waypoints in the other patches of the forest, I was decided to draw the
borders of Kubuh and Mendong II using exclusively the software Google Earth©. The
vegetation composition of these patches could, however, not be estimated.
34
Figure 8: Satellite Photograph of a part of Alegafor with waypoints. Picture taken from
the software Google Earth©.
3.2. Analysing Recorded Signs
3.2.1. Decay Rate
To find and revisit nests, I followed the recorded tracks stored on the GPS device and reached
the recorded waypoints corresponding to them. This aspect of the methodology was biased by
the approximate precision of the GPS device regarding the revisiting of recorded points.
Therefore, the follow-up of recorded nests (which is time consuming) was abandoned, and we
decided to only focus on revisiting “fresh nests”.
In 2008, Kühl et al. presented a range of the average decay rate of chimpanzee nests,
gathering results of numerous previous studies. According to them, the nests of Pan
troglodytes can be visible for more than 220 days (more than seven months). The shortest
decay rate shown in this study is 73 days (over ten weeks). My field work being limited to 3
months, my hopes were to find a fresh nest as early as possible and to revisit it weeks later.
Giving the minimum and maximum of decay rates argued by Kühl et al. (2008), it was
100m
1000 yards
35
necessary to find a fresh nest less than 73 days before my last day on the field, which
corresponds to the 12th
of March. Unfortunately, the first nests were found on March the 18th
,
and the first fresh nest was discovered on the 21st of March. Five other fresh nests spread over
four different nest sites were found afterwards but too close to the end of my study to be
expected to give any practical decay rate (Table 4).
Table 4: Location, date of first observation, and number of days until the end of the
study for each fresh nest collected.
Nest Site Patch
Name Date
Days from the end
of the study
667 Fintse 21/03/2013 64
751 Ndong Efuh 23/04/2013 31
830
Fintse 12/05/2013 12 831
833
With the data collected, no decay rate was expected to be found. Nevertheless, we revisited
the “fresh nest” of the nest site number 667 on the 13th
of May (53 days after its first
observation). Surprisingly, it had completely disappeared. According to these observations,
the decay rate of this nest was strictly inferior to the minimum of 73 days recorded by Kühl et
al. (2008). The same day, another unexpected discovery was made: in the same forest patch, a
very old nest (nest site number 666) which had been discovered on the same date of the fresh
nest from the nest site number 667, was still in the same state of decomposition with clearly
visible branches and a few remaining brown leaves. These observations could be explained by
two factors. First, the tree species on which these two nests were built (and from which the
leaves come from) could be different (both species were unknown). Secondly, the “fresh nest”
was built on a tree located on the edge of an open area with tall grass and opportunistic plants,
while the “old nest” (about 30m away) was situated underneath a closed canopy, in a denser
part of the forest.
The data collected on the chimpanzee nests did not allow me to do any estimation of a global
decay rate. Given that my results show there is a large difference of decay rates between nests
located in the same forest (less than 50m from each other), it is likely that any estimations of
chimpanzee density based on nest density would be highly unreliable (see Discussion).
36
3.2.2. Nest Characteristics
The trigonometric function presented in Figure 5 (see above) was used to determinate the
height of chimpanzee nests based on the measurements recorded in the field.
The cluster size of sleeping groups was recorded using the technique presented by Fruth &
Hohmann (1994): close nests presenting a similar degree of decomposition were described as
belonging to the same nest site.
Chapter 3 – RESULTS
1. Map of Babanki-Finge
The Babanki-Finge forest (Figure 9) is composed of three main ensembles: Alegafor
(belonging to Bambui), Abong-Phen (belonging to Finge and Babanki), and a third set named
Keffem, including the patches surrounding Kedjom-Keku (Fintse, Kubuh, Mendong I, and
Mendong II). It is situated between 1,500 and 2,300m.
1.1. Alegafor
Alegafor is the most Southern patch of the forest of Babanki-Finge, and the closest to
Bamenda (Figure 9). Surrounded by grasslands and farms, and restricted to the valley of a
river (Figure 10), it is a highly disturbed narrow patch. According to Ernest Vunan (one of my
field assistants), Alegafor is a community forest belonging to Bambui. There is no
management over the creation of farms inside the forest, thus the edge of this patch seems to
be continuously nibbled. The anthropomorphic pressure is heavily widespread all over
Alegafor as shown in the map Figure 11.
37
An acacia forest has been planted on the North-West of Alegafor (Figure 10). The species of
acacia planted in this area is not indigenous to the Bamenda Highlands. The reason for the
creation of this artificial forest was to prevent the drought of the stream supplying water to the
village of Bambui.
Kedjom-Keku
Kedjom-
Ketingo
Bambui
Bambili
Sabga
Finge
S
N
W E
Douala
Yaoundé
Bamenda
Finge
Keffem Abong-Phen
Alegafor
1000 km
600 miles
Yaoundé: Political Capital
Douala: Economical Capital
Sabga: Village
: Village with borders
zzAfzefefzefzfincluding Bambafinge
: Patches of the forest
: Stations used during the study
: New built station available
: Borders of Bamenda
: Main road
: Secondary road
: Dirt road
LEGEND:
Figure 9: Map of Babanki-Finge and the towns surrounding the forest.
5 km
3 miles
Bambui
38
Figure 10: Photograph of the middle part of Alegafor. The red circles show the grasslands, the
yellow circle shows the farms (maize) and the blue arrow indicates the valley with the stream. In
the background, the grey circle shows the acacia forest. Credit: Osiris Doumbé.
The area of the totality of Alegafor is about 86ha, with 62% of forest (53ha). Another 18%
(16ha) is covered by disturbed forest with understorey farms. About 5% of the total of
Alegafor (almost 4ha) has been transformed into farms, and over 11% of the patch (around
10ha) exhibits signs of past anthropomorphic pressure. The acacia forest cover is 35.37ha.
1.1. Abong-Phen
The second ensemble of patches is located on the North of the acacia forest of Bambui and is
called Abong-Phen. This area belongs to the villages of Babanki and Finge and includes
several sub-patches: Akwango, Chuh-Nyingong, and Ndong-Efuh.
As Alegafor (but to a lesser degree), Abong-Phen suffers from an uncontrolled
anthropomorphic pressure. In this part of Babanki-Finge, the forest is fragmented by not only
grasslands (Figure 12), but above all by large plantations of garden huckleberry (Solanum
scabrum), locally known as „djama-djama‟, and maize (Figure 13). The map of Abong-Phen
N
39
500m
350 yards
LEGEND:
: Forest
: Riparian Forest
: Disturbed Forest
: Acacia Forest
: Gnidia glauca
: Aframamom sp.
: Fern
: Secondary
amannlyeVegetation
: Shrubs
: Tall Grass
: Farm
: Ancient Farms
: Streams
: Human
JohnnybeGoogd!!!Activities
JohnnybeGogod!!!surrounding
JohnnybeGgood!!!the forest
Grassland
Grassland
Grassland
Grassland
Farms
(maize)
Farms (vegetables)
Farms
(Bananas,
Plantains,
vegetables)
Figure 11: Map of Alegafor and the Planted Acacia Forest
40
is shown in Figure 14. The fragmentation and the extent of this patch are represented in
Figure 15.
Figure 12: Grassland in Southern Ndong-Efuh. Credit: Osiris Doumbé.
Figure 13: Plantation of djama-djama and maize on the edge of the sub-patch of Ndong-
Efuh. Credit: Osiris Doumbé.
41
1875m
1912m
500m
350 yards
CHUH-NYINGONG
AKWANGO
NDONG-EFUH
Grassland
Grassland
Grassland
Farms (eucalyptus &
vegetables)
Farms (maize
& djama-
djama
Farms
(vegetables)
Farms
(djama-
djama)
Farms (bananas,
plantains and
vegetables)
Grassland
Grassland
LEGEND:
: Forest
: Riparian Forest
: Disturbed Forest
: Acacia Forest
: Gnidia glauca
: Aframamom sp.
: Fern
: Secondary
amannlyeVegetation
: Shrubs
: Tall Grass
: Farm
: Ancient Farms
: Streams
: Mounts
: Bare Mountains
: Human
Johnnybeoogd!!!Activities
Johnnybesgod!!!surrounding
Johnnybegood!!!the forest
AKWANGO: sub-patches
Grassland
Figure 14: Map of Abong-Phen
42
Picture 15: Photograph of Abong-Phen. The green polygon encircles Ndong-Efuh, the orange
ellipse shows Akwango, and the two arrows indicate the highest points of Abong-Phen. There is
a clear forest of Gnidia glauca on the left of the foreground. Bamenda is visible in the
background of the photograph. Credit: Osiris Doumbé.
Abong-Phen has an area of 175ha, covered by 108ha of forest. In addition to this 62% of
cover, there is 35% of disturbed forest (61ha) including understorey plantations of coffee.
1.2. Kefem
This third and last set of sub-patches is the most Northern ensemble of Babanki-Finge. It
includes the remaining fragmented forest covers surrounding the village of Kedjom-Keku:
Fintse, Kubuh, Mendong I and Mendong II (Figure 16). Keffem forest is the most difficult to
reach. As an example, Mendong I is more than 5km from Kedjom-Keku, with a total climb of
1000m.
This ensemble is limited in the North by large grasslands. A particularity of this part of the
forest is the cover of bracken ferns (Pteridium aquilinum spp.) on the West and the South of
Mendong I. According to the field assistants Ernest Vunan and „Pa‟ Moses, this whole area
was once farmed, but a conflict between farmers and herders (added to the very long distance
1875m 1918m
N
43
from the village), resulted in the abandonment of the plantations. Bracken ferns are highly
toxic for cattle (Bulnes et al., 2001; Gava et al., 2002), which means this vast area now
remains unexploited.
These sub-patches are divided as family lands according to Ernest. This is why Fintse and
Mendong I (although attached one to another) are considered as two distinctive patches by
villagers. The 17ha and the 113ha of forest cover of respectively Fintse and Mendong I
totalize 73% of the area of the two patches.
The creation of the maps of Kubuh and Mendong II being exclusively made thanks to Google
Earth©, I preferred not to measure the cover of these two areas.
1.3. Comparisons Between Areas
The total forest area of Babanki-Finge is 386.51ha, including 94.13ha of disturbed forest with
understorey farms. These portions of disturbed forest still include a significant number of
mature trees which are left over, in order to cover typical understorey crops such as coco-yam.
The cover areas of the different vegetation types of Alegafor, Abong-Phen, and Mendong I-
Fintse are presented in Table 5. Amongst the three ensembles of patches, Alegafor appears to
be the most disturbed patch, while Mendong I and Fintse are shown as possessing the largest
forest cover. Table 6 shows the covers of present and former human sedentary disturbances
(i.e. agriculture or grazing) on the forest. This table shows Mendong I and Fintse seem to
undergo less anthropomorphic sedentary pressure nowadays than in the past. On the contrary,
farming seems to be increasing in the other two ensembles, and particularly in Abong-Phen.
Aframomum sp. is commonly found concentrated in open-areas and on the edge of the forest.
It is also generally dominant in the understorey of the forest. The fruits of these plants are
widely eaten by monkeys (Agmen et al., 2010), apes (Wrangham et al., 1994), and humans
(pers. obs.). Because most of the open areas dominated by Aframomum were located on the
edge of the forest, I concluded they were signs of past human disturbances (i.e. farming or
burning). Gnidia glauca trees were also used as signs of past human disturbances as they are
typical of secondary forests, and appear to be the transition between degraded and mature
forests (Ingram & Nsom Jam, 2007).
44
LEGEND:
: Forest
: Riparian Forest
: Disturbed Forest
: Acacia Forest
: Gnidia glauca
: Aframamom sp.
: Fern
: Secondary
amannlyeVegetation
: Shrubs
: Tall Grass
: Farm
: Ancient Farms
: Streams
: Mounts
: Bare Mountains
: Human
JohnnybeGogd!Activities
JohnnybeGgod!surrounding
Johnnybegood!the forest
KUBUH: sub-patches
Grassland
KUBUH
MENDONG I
MENDONG II
FINTSE
Farms (sweet
potatoes, maize &
cassava)
Grassland
Grassland
Unexploited
land covered
by ferns
Unexploited
land covered
by ferns
500m
0.2 mile
2292m
2214m
Figure 16: Map of Kefem
45
In all patches, the forest is almost exclusively in valleys along streams. Therefore, the slope
was mostly steep or precipitous. The flattest area is situated in the East of Abong-Phen, but is
gradually transformed into disturbed forests.
Table 5: Cover areas of the different vegetation types encountered in the three
ensembles measured. *: the forest cover includes the cover of the riparian forest.
Vegetation
Description
Alegafor Abong-Phen Mendong I & Fintse
ha % ha % ha %
Forest* 53.73 62.19 108.00 62.14 130.65 73.50
Disturbed
Forest
16.18 18.72 61.34 35.03 16.61 9.34
Farm 4.82 5.58 3.41 1.95 1.17 0.66
Aframomum sp. 1.73 2.00 0.15 0.08 0.22 0.12
Gnidia glauca 0.13 0.15 0.87 0.49 4.74 2.67
Fern 3.61 4.18 0.00 0.0 14.60 8.21
Secondary
Vegetation
2.98 3.45 0.13 0.07 0.40 0.22
Shrubs 0.28 0.32 0.00 0.00 0.00 0.00
Tall Grass 1.36 1.58 0.00 0.00 9.30 5.23
Ancient Farms 1.54 1.78 0.35 0.20 0.00 0.00
TOTAL 86.40 170.28 177.74
Table 6: Cover of farms and areas with signs of former human sedentary pressure. In
this table, the category disturbed forest is included in the farms. „Former human
sedentary pressure‟ gathers ancient farms, and areas with non-climax habitat: Gnidia
glauca forests, Aframomum sp., ferns, secondary vegetation, shrubs, and tall grass.
Habitat Alegafor Abong-Phen Mendong I & Fintse
Farms 24.30% 36.98% 10.01%
Former Human
pressure 12.98% 1.03% 16.74%
2. Observations and Signs of Monkeys and Apes
2.1. Direct Signs
Two species of monkeys were directly observed: the putty-nosed monkey (Cercopithecus
nictitans martini) and the tantalus monkey (Chlorocebus tantalus tantalus) respectively
locally named “black monkey” and “white monkey”. All the direct signs of monkeys were
46
made in the Fintse patch. The direct observation of the putty-nosed monkey was preceded by
alarm calls of at least two individuals. One mature monkey was seen but the noise and
movements observed in the trees around it suggested at least three other individuals. Two
other mature Cercopithecus nictitans martini were found killed by a hunter. The observed
tantalus monkey was caught in a trap and was a juvenile. During our field work in Mendong I,
another hunter stated he caught a mother Chlorocebus tantalus tantalus with its infant.
According to local villagers, putty-nosed monkeys and tantalus monkeys are the two most
observed and hunted primates in the area. Two other species of monkeys were described by
monitors and farmers as occurring in small numbers in the area: the baboon (Papio anubis),
and the locally named “red monkey” which could refer to the patas monkey (Erythrocebus
patas). None of these species was observed during the field work. Although signs of
chimpanzees were found (see below) they were never directly observed.
The local villagers argued monkeys tend to come to their plantations to feed on maize during
the harvesting season (July-August). The field assistant David Achomochi also commented,
stating tantalus monkeys are also locally well-known for destroying coco-yam plantations. On
the contrary, around Babanki-Finge, chimpanzees are less seen as threats by farmers, rarely
crop raiding. Nevertheless, several stories of dog-killer chimpanzees have been heard. These
anecdotes would have happened when domesticated dogs (Canis lupus familiaris) try to chase
chimpanzees. Therefore, villagers specified the apes do not kill dogs for their meat, but to
defend themselves.
2.2. Indirect Signs
2.2.1. Chimpanzee Nests
The indirect signs of haplorhines were mostly chimpanzee nests (Figure 17). A total of 51
nests were found during my field work in Babanki-Finge. Another 20 nests were discovered
by my field assistants who pursued the data collection once I left Cameroon. The
characteristics of each nest are in Table 7.
The distribution of the sleeping sites is presented in Figure 18 and Figure 19. Most of the
nests were found in the Northern part of the forest (52 nests) and none were discovered in
Alegafor. Amongst the fragmented sub-patches of Abong-Phen, chimpanzee nests were only
found in Ndong-Efuh (19 nests). All the nests were considered as sleeping nests because there
47
was no ways to differentiate night to day nests. Therefore, all the nesting sites found were
expected to be sleeping sites.
Figure 17: Old nest of Chimpanzee in the canopy. Credit: Ernest Vunan.
Table 7: Characteristics of the Recorded nests. Unknown species of trees found are
noted Ukn. *: the nests found in Kubuh and Mendong II were recorded by the monitors,
therefore height and sizes were not measured nor estimated.
Date Patch Age Height
(m) Tree Species #
18/03
Mendong I
very old
9.39 Carapa grandiflora
1
Old 2
Old 3
Old 4
Old 9.01 Carapa grandiflora 5
old 7.83 Carapa grandiflora
6
old 7
old 8.52 Carapa grandiflora 8
old Carapa grandiflora 9
16/04 very old 19.76 Carapa grandiflora 10
17/04 old Ukn 11
48
Table 7 (following): Characteristics of the Recorded nests. Unknown species of trees
found are noted Ukn. *: the nests found in Kubuh and Mendong II were recorded by the
monitors, therefore height and sizes were not measured nor estimated.
Date Patch Age Height
(m) Tree Species #
21/03
Fintse
old 10.75 Ukn 12
old 7.92 Carapa grandiflora 13
old 6.87 Carapa grandiflora 14
old 5.77 Carapa grandiflora 15
recent Ukn
16
very old 17
old 10.49
Carapa grandiflora 18
old Carapa grandiflora 19
very old 13.55 Ukn 20
fresh Ukn 21
12/05
old 8.5 Strombosia sp. 22
fresh 6.39 Carapa grandiflora 23
fresh 7.48 Strombosia sp. 24
old 6.44 Strombosia sp. 25
old 6.82 Carapa grandiflora
(and Strombosia sp.)
26
fresh (of the
night)
Carapa grandiflora
(and Strombosia sp.)
27
old Carapa grandiflora 28
old Strombosia sp. 29
fresh
Carapa grandiflora
30
old 31
very old 32
08/04
Abong-Phen
old 11.94 Ukn 33
22/04
very old 18.54 Carapa grandiflora 34
recent 8.13 Carapa grandiflora 35
recent 14.70 Ukn 36
old 8.31 Carapa grandiflora 37
old 10.54 Carapa grandiflora 38
old 7.52 Carapa grandiflora
39
recent 6.90 40
old 7.24
Carapa grandiflora
41
old 7.71 42
old 9.25 43
23/04
very old 11.88 Strombosia sp. 44
fresh 8.42 Ukn 45
recent 6.43 Ukn 46
very old 8.42 Ukn 47
old 4.99 Ukn 48
49
Table 7 (following): Characteristics of the Recorded nests. Unknown species of trees
found are noted Ukn. *: the nests found in Kubuh and Mendong II were recorded by the
monitors, therefore height and sizes were not measured nor estimated.
Date Patch Age Height
(m) Tree Species #
06/05 Abong-Phen
old 12.46 Carapa grandiflora 49
old 11.59 Carapa grandiflora 50
09/05 old 6.97 Ukn 51
28/05 Kubuh*
old Strombosia sp. 52
old Strombosia sp. 53
old Strombosia sp. 54
old Ukn 55
03/06
Mendong II*
old Carapa grandiflora 56
old Carapa grandiflora 57
old Carapa grandiflora 58
old Carapa grandiflora 59
old Carapa grandiflora 60
old Strombosia sp. 61
old Carapa grandiflora 62
old Carapa grandiflora 63
old Carapa grandiflora 64
04/06
old Carapa grandiflora 65
old Carapa grandiflora 66
old Carapa grandiflora 67
old Carapa grandiflora 68
old Carapa grandiflora 69
old Carapa grandiflora 70
old Carapa grandiflora 71
During the data collection made by Ernest and the monitors in June, the patch of Mendong II
was described as rich in Ficus, Carapa and Strombrosia trees, as well as rich in water
streams.
Chimpanzee Nests - Tree Species
The lack of an extensive guide of the trees of the Babanki-Finge forest prevented us from
recording all the species of trees in which chimpanzee nests were found. Only two species of
trees were identified (Table 8). The unknown species found were composed of at least four
different taxa. The percentages of the species of trees used by chimpanzees for nesting are
shown in Figure 20.
50
1875m
1912m
500m
350 yards
CHUH-NYINGONG
AKWANGO
NDONG-EFUH
Grassland
Grassland
Grassland
Farms (eucalyptus &
vegetables)
Farms (maize
& djama-
djama
Farms
(vegetables)
Farms
(djama-
djama)
Farms (bananas,
plantains and
vegetables)
Grassland
Grassland
LEGEND:
: Forest
: Riparian Forest
: Disturbed Forest
: Acacia Forest
: Gnidia glauca
: Aframamom sp.
: Fern
: Secondary
mannlyeVegetation
: Shrubs
: Tall Grass
: Farm
: Ancient Farms
: Streams
: Mounts
: Bare Mountains
: Human
JohnnybeGood!!!Activities
JohnnybeGood!!!surrounding
JohnnybeGgod!!!the forest
AKWANGO: sub-patches
: nesting sites
Grassland
Figure 18: Map of the chimpanzee nesting sites in Abong-Phen
51
LEGEND:
: Forest
: Riparian Forest
: Disturbed Forest
: Acacia Forest
: Gnidia glauca
: Aframamom sp.
: Fern
: Secondary
mannlyeVegetation
: Shrubs
: Tall Grass
: Farm
: Ancient Farms
: Streams
: Mounts
: Bare Mountains
: Human
ohnnybeGoogd!Activities
ohnnybeGogod!surrounding
ohnnybeGgood!the forest
KUBUH: sub-patches
: nesting sites
Grassland
KUBUH
MENDONG I
MENDONG II
FINTSE
Farms (sweet
potatoes, maize &
cassava)
Grassland
Grassland
Unexploited
land covered
by ferns
Unexploited
land covered
by ferns
500m
0.2 mile
2292m
2214m
Figure 19: Map of the chimpanzee nesting sites in Kefem
52
All nests but two were built in single trees. The nests built in two trees used mixed species:
Strombosia sp. and Carapa grandiflora.
Table 8: Species of trees in which chimpanzee nests were found. Ukn=Unknown taxa.
Species of Tree Number
of Nests
Carapa grandiflora 46
Strombosia sp. 9
Ukn 16
Carapa grandiflora and Strombosia sp. 2
Figure 20: Exploded pie representing the species of trees used by the chimpanzees of
Babanki-Finge for nesting. Percentages are rounded up.
Chimpanzee Nests - Height
The height was not recorded for all nests because of clinometer problems in very bright areas.
Out of the 51 nests recorded during my field work, the height of 40 of them was measured.
The descriptive statistics are shown in Table 9 and the distribution of the nests in the layers of
Carapa grandiflora
65% Strombosia sp.
12%
Carapa g. & Strombosia sp.
3%
Unknown 20%
Tree Species Used by the Chimpanzees for Nesting
53
the forest is shown in Figure 21. According to these results, most of the recorded nests were
found in the small layer of the forest (below 11m). None of the nests have been found on the
ground.
Table 9: Descriptive Statistics of the measured height of 40 nests (in meters).
Mean Median Minimum Maximum
9.12 (sd=3.08) 8.46 4.99 19.76
Figure 21: Distribution of the chimpanzee nests in the layers of the trees of the forest.
Chimpanzee Nests - Cluster Size
Nests were found in very small groups. Cluster size of nesting groups ranged from 1 to 4, with
an average of 1.39 nests (sd=0.68, n=71). The numbers of nests per sleeping site are shown in
Table 10 and illustrated in the graph of Figure 22.
Chimpanzee nests of the same nesting site were almost always found in individual trees. Only
two sleeping sites (both in Mendong I) showed sleeping nests sharing the same tree. In one
nesting site, one tree was shared by three nests, and in another site, two nests were built on the
same tree.
2.5% (1/40)
77,5% (31/40)
20% (8/40)
0% (0/40)
0 10 20 30 40 50 60 70 80 90
0 to 4.99m
5.00 to 10.99m
11.00 to 29.99m
over 30m
Percentage of Nests
He
igh
t o
f th
e N
est
Nest Heights
54
Table 10: Number of Nests per Sleeping Site.
Number of Nests
per Sleeping Site
Number of
Sleeping Sites
1 36
2 11
3 3
4 1
Figure 22: Frequency of chimpanzee nest group sizes in Babanki-Finge.
Chimpanzee Nests - Nesting Site Choice
All nests but one were located in the forest. The remaining nest was found in the disturbed
forest of Ndong-Efuh (Abong-Phen). Out of the 27 nesting sites found, 20 were located under
a closed canopy and seven under a canopy of 51-75% coverage. The chimpanzees of Babanki-
Finge seem to prefer nesting in areas with dense vegetation.
70.59%
21.57%
5.88% 1.96%
0
10
20
30
40
50
60
70
80
1 2 3 4
Fre
qu
en
cy o
f th
e s
ize
of
Sle
ep
ing
S
ite
s
Group Size
Nests Per Group
55
2.2.2. Other Signs
Other than nests, one chimpanzee track and two chimpanzee footprints were found in
Northern Mendong I.
Chapter 4 - DISCUSSION
This part of the thesis starts by examining the results, discussing the created habitat maps, and
analysing the anthropomorphic pressure which can be assessed from them. Secondly, the
density, measurements and characteristics of the collected nests are reviewed and compared
with previous studies from other sites of Africa. Finally, some conservation actions will be
proposed, followed by suggestions for further studies in the Babanki-Finge forest and more
generally in the North-West region of Cameroon.
1. Forest Mapping
Babanki-Finge is composed of three sets of patches (Figure 23). Fragmented throughout an
uneven landscape, the remaining forest cover of the area is over 440. Like the pockets of the
forest neighbouring Budongo and Kibale National Park (Uganda) described by McLennan
(2008) and Chapman et al. (2003), Babanki-Finge has been modified for years by
anthropomorphic activities (deforestation, clearing, farming) and is now mostly restricted to
valleys and watercourses.
56
LEGEND:
: Forest
: Riparian Forest
: Disturbed Forest
: Acacia Forest
: Gnidia glauca
: Aframamom sp.
: Fern
: Secondary
mannlyeVegetation
: Shrubs
: Tall Grass
: Farm
: Ancient Farms
: Streams
: Mounts
: Bare Mountains
500m
350 yards
500m
350 yards
Figure 23: Map of the forest patches of Babanki-Finge.
650m
450 yards
5 km
3 miles
57
1.1. Characteristics of the Vegetation
1.1.1. Forests
The forest cover of Babanki/Finge is mainly concentrated in valleys, following streams. It is
dominated by mature trees such as Carapa grandiflora, Ficus sp., and Strombosia sp. It is
highly fragmented and distributed as pockets throughout the area. Abong-Phen is the most
fragmented ensemble of patches, but also the part of Babanki-Finge with the most important
forest cover: 169.34ha (in comparison to Mendong I & Fintse, and Alegafor which
respectively possess 147.26ha and 69.91ha of forest).
Typically, a riparian forest defines the woodland bordering the banks of a stream (Doumbé,
2011). For the study site of Babanki-Finge, because most of the forest cover is surrounding
streams, I decided to specify this habitat as the peculiar climax vegetation alongside some
parts of the water, dominated by an unknown tree species covered with moss and Cyathea
manniana. These medium-height plants create a darker and more humid understorey (pers.
obs.) than in the nearby non-riparian forest.
In Northern latitudes, riparian forests are known for playing an important role of nursery for
aquatic animals (Doi, 2009) and for breeding birds (Darveau et al., 1995). In tropical islands,
where most of fishes and Crustaceans are diadromous amphipods (Monti et al., 2010), borders
of streams are not used for nurseries, but a correlation between shrimps (Macrobrachium sp.
and Xiphocaris sp.) and litter (brought by bordering trees) is expected (Doumbé, 2011). In
Babanki-Finge, the role of this specific forest is still unknown.
1.1.2. Aframomum sp.
About 50 species of rhizomal herbs from the genus Aframomum (Zingiberaceae) occur in
West and central Africa (Tane et al., 2005). The species of the widespread Aframomum of
Babanki-Finge was not determined in this study. It is largely used by haplorhines, including
humans, and seed dispersal by these species may explain its wide distribution throughout the
site of Babanki-Finge. It is a tall grass which produces a red fruit with a fleshy pulp (Figure
24). The seeds of this plant have been reported to be the most abundant in faecal samples of
Chlorocebus tantalus tantalus collected at Nyel Ngaki (Nigeria) by Agmen et al. (2009).
Aframomum sp. fruits have also been observed to be eaten by humans (pers. obs.) and Pan
troglodytes (Tutin & Fernandez, 1997; Wrangham et al., 1994), providing an extensive seed
58
dispersal both throughout the understorey and the open areas of the forest. In addition, several
species of Aframomum are used in medicine and in the ethnodietary preparation of traditional
dishes: the main ingredient of the sauce of the “Mbongo Tchobi” of the Basaa tribe of the
Littoral region of Cameroon is Aframomum citratum (Tane et al., 2005).
Figure 24: Photograph of fruits and leaf of Aframomum sp. Credit: Osiris Doumbé.
In addition to its consumption, Aframomum sp. is also used as tools in certain populations of
chimpanzees (Pan troglodytes spp.). Hashimoto et al. (2000) have reported the straightness
and flexibility of the stem of this tall grass to be particularly appreciated and used as tools by
ant-dipping chimpanzee groups in the Kalinzu forest (Uganda). It is unknown whether the
chimpanzees of Babanki-Finge use this plant for the same purpose or not.
59
1.1.3. Grassland
The forest patches of Babanki-Finge are separated by grasslands. This clear-cut vegetation is
the origin of the given name “Grassfields” to the mountains of the North-West region of
Cameroon. The high altitude of the Bamenda Highlands (and the Adamawa mountainous
massif) provides an environment tse-tse fly free, particularly important for Mbororo
pastoralists (Boutrais, 1992, Youta Hapi, 1998).
1.2. Human Impact
The anthropomorphic pressure has resulted in extensive fragmentation of the dense
vegetation, restricting the remaining forest patches to areas that are relatively inaccessible to
people, such as valleys and precipitous slopes of high mountains.
Table 11 presents the percentage of areas with signs of ancient human clearings, and areas
with current farms, for each ensemble of patches. This table shows Abong-Phen has the
largest percentage of farms (36.98%), and Mendong I and Fintse the biggest percentage of
abandoned clearings (16.35%). These results corroborate the general tendency described by
my field assistant Ernes Vunan. According to him, because of the former conflicts with
powerful herders and the very long distance to reach Mendong I and Fintse, farmers from
Babanki now prefer to create new plantations in Abong-Phen, that are easier to reach from
heir homes. Therefore, farms in Mendong I and Fintse are more and more abandoned and
conquered by opportunistic plants, while the forest of Abong-Phen is more and more cleared
for the production of djama-djama and maize. This observed tendency of focusing in some
patches instead of others is possible for Babanki people because most of the forest belongs to
them, but not for Bambui people who only own Alegafor and therefore target this pocket
particularly. Finge people can only farm in Abong-Phen, but they are far less numerous than
Babanki or Bambui people and therefore exercise a lighter pressure.
The abandoned farms are quickly recovered by opportunistic plants (Aframomuum sp., tall
grass, fern) or gradually conquered by the forest (Gnidia glauca).
60
Table 11: Cover areas of the different vegetation types encountered in the three
ensembles measured. „Farms‟ category includes: Farms and Disturbed Forest; „Former
human sedentary pressure‟ category includes: Ancient Farms, Gnidia glauca Forests,
Ferns, Secondary Vegetation, Shrubs, and Tall Grass.
Habitat Alegafor Abong-Phen Mendong I & Fintse
Farms 24.30% 36.98% 10.01%
Former Human
pressure 12.98% 1.03% 16.74%
The human pressure is uneven throughout Babanki-Finge. As a consequence, even though
monkeys and apes seem to be absent in the most Southern patch (Alegafor), some diurnal
primates still inhabit other areas.
2. Pan troglodytes ellioti: Discussion and Comparisons
In this part of the discussion, many study sites with Pan sp. will be mentioned and compared
to the results found in the with the population of chimpanzees of Babanki-Finge. Figure 25
presents the location of these sites throughout Africa. Some bonobo (Pan paniscus) sites are
included in the comparisons as this species has many similarities with Pan troglodytes,
although there are also many social and ecological differences between the two species of Pan
(Chapman et al., 1994).
2.1. Chimpanzee Density
The study failed to provide an estimation of the population of Pan troglodytes ellioti of
Babanki-Finge for a number of reasons. Firstly, the methodology was based on nest counting
which needs to find fresh nests as early as possible. Because the first targeted area was
Alegafor (where no signs of chimpanzees were found), we did not succeed in finding a nest
before mid-March. Then, the length of the field work was too short to follow-up a sufficient
enough number of fresh nests. The shortness of the length data collection provided a second
flaw to this study: in just three months, the inter-annual variation in density of chimpanzee
populations argued by Tutin & Fernandez (1984) cannot be taken into account.
61
Figure 25: Map of the sites mentioned in this section.
LEGEND
: Study Site of this thesis.
Study Sites with Pan troglodytes verus: 1: Niokolo-Koba NP (Baldwin et al., 1981; Fruth &
Hohmann, 1994), 2: Bossou (Matsuzawa & Yamakoshi, 1996; Sugiyama, 1998), 4: Nimba
mountains (Matsuzawa & Yamakoshi, 1996), 4: Sapo NP (Fruth & Hohmann, 1994), 5: Taï NP
(Fleury-Brugière & Brugière, 2010; Fruth & Hohmann, 1994; Kouakou et al., 2009).
Study Site with the unidentified subspecies of Pan troglodytes sp.: 6: Ise FR (Ogunjemite et
al., 2006).
Study Sites with Pan troglodytes ellioti: 7: Gashaka-Gumti NP (Sommer et al., 2004), 8:
Takamanda NP (Sunderland-Groves et al., 2003), 9: Bechati-Andu (Last & Muh, 2013), 10:
Banyang-Mbo Sanctuary (Greengrass & Maisels, 2007).
Study Sites with Pan troglodytes troglodytes: 11: Campo NP & Ma’an Forest (Matthews &
Matthews, 2004), 12: Ntonga (Dupain et al., 2004), 13: Dzanga-Ndoki NP (Blom et al., 2001), 14:
Lopé NP (Fruth & Hohmann, 1994; Tutin et al., 1997; Tutin, 1999), 15: Odzala NP (including
Lokoué) (Devos et al., 2008), 16: Lake Télé (Poulsen & Clark, 2004), 17: Nouabalé-Ndoki NP
(including Goualougo Triangle) (Devos et al., 2008).
Study sites with Pan troglodytes schweinfurthii: 18: Budongo (Chapman et al., 2003; Fruth &
Hohmann, 1994; Hashimoto, 1995; McLennan, 2008; Plumpre & Reynolds, 1997; Tutin &
Fernandez, 1984), 19: Kibale NP (including Ngogo and Kanyawara) (Chapman et al., 2003;
Hashimoto, 1995; Tutin & Fernandez, 1984), 20: Virunga NP (Fruth & Hohmann, 1994), 21 Lake
Kivu (Fruth & Hohmann, 1994), 22: Kahuzi-Biega NP (Basabose & Yamagiwa, 2002; Hall et al.,
1998), 23: Gombe NP (Fruth & Hohmann, 1994), 24: Ugalla GR (Fruth & Hohmann, 1994;
Ogawa et al., 2007).
Study sites with Pan paniscus. 25: Lake Tumba (Fruth & Hohmann, 1994), 26: Lomako NP
(Fruth & Hohmann, 1994), 27: Wamba (Fruth & Hohmann, 1994), 28: Yalosidi (Fruth &
Hohmann, 1994), 29: Salonga NP (including Lokofa, Etate, Beminyo, Isanga,Yongo and Ikolo)
(Fruth & Hohmann, 1994).
FR: Forest Reserve; GR: Game Reserve; NP: National Park
20
21
9 4 5
6 Babanki-Finge
1
2 3
8 7
11 13
14 25
26
27
28 29
18 19
22
2
23
2 24
2
Babanki-Finge
12
16
17 15
10
62
Another difficulty encountered during the data collection was the character of the field.
Although the forest is small, its fragmentation and the ruggedness of its terrain make it hard to
cover. Because we stayed in each patch for several days, in a group of up to five individuals
(including the two monitors, Ernest, a photographer, and myself), slowly covering the forest,
it is much probable that chimpanzees and monkeys fled before any observation. It would also
explain why we found so few fresh nests.
Finally, the data collection did not allow us to assess if a nest was used more than once.
Nevertheless, the reuse of night nests is not rare (10.8% nests are reused), particularly for
injured chimpanzees which have been reported to reuse 20.7% of their nests (Plumptre &
Reynolds, 1997).
Even though not hunted, the chimpanzees of Babanki-Finge are not expected to be numerous
regarding the number of collected nests and the degree of fragmentation of the area. Other
studies conducted in small forest patches have suggested an average group size of 12-20
individuals (McLennan, 2008). The largest number of similar aged nests found together in
Babanki-Finge was four, suggesting that this is the minimum number of chimpanzees living
in the forest.
2.1.1. Nest Counting
Nest counting has been described as the best method to estimate the population of great apes
over large areas, due to the low density of gorillas and chimpanzees (Tutin & Fernandez,
1984). Nevertheless, it is also well-known for its limitations (Plumptre & Reynolds, 1996).
Accuracy of Decay Rate
Assessing a decay rate is essential to estimate the density of an ape population based on nest
counting (White & Edwards, 2000). However, decay rates of chimpanzee nests are highly
dependent on seasons and sites (Dupain et al., 2004; Last & Muh, 2013; Hall et al., 1998).
Therefore, it is preferred to calculate the nest decay rate for each site, which can be tricky
because of the need to follow-up a significant number of fresh nests (Tutin & Fernandez
1984).
63
In this study, the results have failed to assess a decay rate because of the lack of significant
data on fresh nests. Nevertheless, the collected data from Babanki-Finge have shown the
estimation of a decay rate must be taken extremely carefully because of the differences in
vegetation cover within a study site. A comparison of two nests near to one another illustrates
the problem of estimating decay time. On the 21st of March, a fresh nest (nest site 667) and a
very old nest (nest site 666) were found 30m apart. During the re-visit of the area (53 days
after the discovery of the nests), while „very old‟ nest 666 was found in the same shape as
previously, „fresh‟ nest 667 had completely disappeared.
The range of decay rates within a single site is usually very extensive. In a Gabonese site, out
of 49 nests the decay rate of Pan troglodytes troglodytes was measured to range from 35 to
151 days (Tutin & Fernandez, 1984), while in Taï National Park (Ivory Coast), the average
decay rate was estimated at 62.92 days, with a standard deviation of 51.79 days (Kouakou et
al., 2009).
Commonly used methods of calculations of great ape density from nest counts include the use
of:
Equation 1 (Tutin & Fernandez, 1984):
=
( ) × ( )
Equation 2 (Hashimoto, 1995):
=
×
×
× [ ]
×
Nevertheless, because of their position up in trees, chimpanzee nests (probably more than
gorilla nests which are mostly built on the ground) tend to show a very high variability in
64
decay rates. The canopy protects nests more or less against the climatic conditions (strong
winds or heavy rain). Therefore, the decay rate of a chimpanzee nest might strongly depend
on the cover of the vegetation surrounding the tree where it is built. Unfortunately neither the
equation of Tutin & Fernandez (1984), or the improved equation of Hashimoto (above) take
this factor into account. As it was pointed out by Sanz et al. (2007), it is urgent to increase the
accuracy of the methodology of chimpanzee and gorilla surveys.
Even though nest counting may give a good estimation of group sizes and of relative
population sizes of great apes, it should not be used as an accurate method to calculate gorilla
or chimpanzee density. Above all, this method should not be used as a mean to compare
populations from different areas, or new studies to older studies from the same site. The
accuracy of the decay rate, maybe even more than environmental factors, might be the reason
why surveys on same sites give different ape densities (Table 12).
Table 12: Comparisons of density estimations from previous studies.
Taxon COUNTRY Study
Site Author Density
Pan
troglodytes
verus
Ivory Coast Taï
Kouakou et
al., 2009
0.69-
1.75
Fleury-
Brugière &
Brugière, 2010
0.10
Pan
troglodytes
schweinfurthii
Uganda
Kibale
Hashimoto,
1995 1.97
Tutin &
Fernandez,
1984
2.5-3.8
Budongo
Hashimoto,
1995 1.3-2.3
Tutin &
Fernandez,
1984
6.7
Because of its relative simplicity and advantaging cost, the nest counting method is widely
used. It should however be restricted to measure and compare nest densities rather than
chimpanzee densities like it has been discussed by Plumptre & Reynolds (1997).
65
2.1.2. Nest Density
Chimpanzees seemed to find sufficient requirements for nesting in Fintse (123 nests/km²)
despite its small area (17ha) (Table 13). According to the report of my field assistant Ernest
Vunan after their data collection in Mendong II, the patch presented a lot of Capara
grandiflora, Strombosia sp. and Ficus sp. as long as watercourses. This noted abundance in
trees used by chimpanzees explained the discovery of 16 nests in the patch.
The density of nests per km² was expected to be higher in the study site than in other
chimpanzee locations because the limits of the home range of the population of Pan
troglodytes ellioti of Babanki-Finge are unknown. Moreover, the study focused on a small
area due to the shortness of the data collection, and the lack of pilot studies. In comparison to
other chimpanzee sleeping sites (Table 13) the density of Pan troglodytes ellioti found in
Babanki-Finge is quite high: 17 nests/km in the Abong-Phen-Fintse-Mendong I area, against
1.0 and 7.2 nests/km² in Lokoué and Gouoalagou Triangle (Congo). This high difference of
density is mostly due to the fragmentation of the forest of Babanki-Finge, providing fewer
nesting trees than in the two Congolese sites. Moreover, the total area of Abong-Phen-Fintse-
Mendong I is far smaller than the areas to which it is compared to.
The density of chimpanzees in the unprotected forest of Ntonga (49.9 nests/km²) was
described as surprisingly higher than in the neighbouring Dja faunal reserve (Dupain et al.,
2004), while the average of 33.3 nests/km² in Banyang-Mbo is explained by the fact it is one
of the healthiest population of Nigeria-Cameroon chimpanzees (Morgan et al., 2011).
2.2. Nest Height
The differences in nest height are bigger within a population than between populations of Pan
sp., but also significantly more important between bonobos and chimpanzees (Fruth &
Hohmann, 1994) as shown in Table 14. Pan paniscus seem to build nests higher than Pan
troglodytes.
Out of the 13 means of height presented in Table 14, Babanki-Finge (Cameroon) and Kahuzi-
Biega (DRC) are the sites where chimpanzees build nests the lowest on average (respectively
9.12m and 9.4-9.8m). The explanation of this particularity is probably the altitude of the two
sites. With 1500-2300m and 2050-2350m of altitudes, Babanki-Finge and Kahuzi-Biega are
66
Table 13: Comparison of chimpanzee nest densities between some patches of Babanki-
Finge and other sites. The habitat characteristic codes are the followings: GWF: mosaic
of Grassland and Woodland Forest; RF: RainForest.1: located in the Odzala National
Park; 2: located in the Nouabalé-Ndoki National Park.
Subspecies COUNTRY Study
Site Author
Habitat
&
Altitude
Area
(km²)
Number
of Nests
Nest
Density
(nests/km²)
Pan
troglodytes
elloti Cameroon
Abong-Phen
Doumbé,
2013
GWF
(1500-
2300m)
1.70 19 11.15
Fintse 0.17 21 123.52
Mendong I 1.13 11 9.73
Abong-Phen, Fintse
and Mendong I 3 51 17
Banyang-Mbo
Greengrass
& Maisels,
2007
RF
(120-
1750m)
660 - 33.3
Pan
troglodytes
troglodytes
Ntonga Dupain et
al., 2004
RF
(<1000m) 30 247 49.9
Congo
Lokoué1
Devos et
al., 2008
RF
(300-
600m)
42 - 1.0
Goualougo Triangle2
RF
(<1000m) 380 - 7.2
by far the highest sites presented in this table. In DRC, the forest mean height of trees in the
mountainous forests of DRC (1730-2820m) was averaged at 19.81m, against 29.87m in
forests below 1200m (Grubb et al., 1963). The general tendency of trees to shorten with
altitude probably also applies to nesting trees, which would explain the lower height of nests.
Males have also been reported as nesting in lower layers than females (Brownlow et al.,
2001). Unfortunately, we are unable to determine from our data whether the gender affected
the height of nests in Babanki-Finge or not.
The maximum height of the nests of Babanki-Finge is one of the smallest of Table 14
(19.79m). However, the minimum nest height is the highest amongst the mentioned sites: the
lowest nest was found at 4.99m. Unlike in several other sites, no ground nest was found in
Babanki-Finge.
Terrestrial Nests
In addition to arboreal nests, chimpanzee sleeping sites can also contain terrestrial nest.
Matsuzawa & Yamakoshi (1996) have argued the weaned individuals of the Pan troglodytes
67
Table 14: Comparison of nest heights between Babanki-Finge and populations of Pan
troglodytes spp. and Pan paniscus from other sites. The habitat characteristic codes are
the followings: GWF: mosaic of Grassland and Woodland Forest; RF: RainForest; SW:
Savannah-Woodland. *: even though 71 nests were found in this study, the height of
only 40 was measured. (?): unidentified subspecies of chimpanzee from South-Western
Nigeria.
Subspecies COUNTRY Study
Site Author
Habitat &
Altitude
Number
of Nests
Height (m)
Mean
(sd) Median Range
Pan
troglodytes
verus
Senegal Niokolo-
Koba
Baldwin et
al., 1981 SW
(<500m)
252 - 11 0-44
Fruth &
Hohmann,
1994
4,478 - - 5-22
Republic
of Guinea -
SW 100 11.5 - 4-
31.5
- 184 - - 2-24
Liberia Sapo SW
(<500m) 67 - 12 6-20
Ivory
Coast Taï
RF
(<500m)
146 - - 3.5-
15
154 23.2 20 5-45
Pan
trogoldytes
spp. (?)
Nigeria Ise Ogunjemite
et al., 2006
RF
(<500m) 61
19.14
(6.75) - -
Pan
troglodytes
Ellioti
Cameroon
Bechati-
Andu
Last &
Muh, 2013
RF
(200m-
1700m)
113 - - 0->40
Bamenda
Highlands
Doumbé,
2013
GWF
(1500-
2300m)
40* 9.12
(3.08) 8.46
4.99-
19.79
Pan
troglodytes
troglodytes
Equatorial
Guinea -
Baldwin et
al., 1981 RF 195 - 10 0-20
Fruth &
Hohmann,
1994
195 - 10 0-40
Gabon Lopé RF
(<1000m)
1,741 8.7 - 2-32
523 11.7 10 2-45
Pan
troglodytes
schweifurthii
Uganda
Budongo
Plumptre &
Reynolds,
1997 RF
(~1,100m)
374 - -
From
0-5 to
31-35
Fruth &
Hohmann,
1994
259 - - 3-45
Ngogo RF
372 12.2 - 2-35
Kanyawara 63 10.3 - 5-23
Tanzania Gombe
GWF 384 - - 12-24
Ugalla 491 19 - 5-40
DRC Lake Kivu RF - - - 10-25
Virunga GWF 101 13.5 - -
68
Table 14 (following): Comparison of nest heights between Babanki-Finge and
populations of Pan troglodytes spp. and Pan paniscus from other sites. The habitat
characteristic codes are the followings: GWF: mosaic of Grassland and Woodland
Forest; GRF: mosaic of Grassland and RainForest; RF: RainForest; SW: Savannah-
Woodland.
Subspecies COUNTRY Study
Site Author
Habitat &
Altitude
Number
of Nests
Height (m)
Mean
(sd) Median Range
Pan
troglodytes
schweifurthii
DRC
Kahuzi-
Biega
Basabose
&
Yamagiwa,
2002
RF/Primary
Forest
(2050-
2350m)
104 9.4 - -
RF/Secondary
Forest
(2050-
2350m)
94 9.8 - -
Pan paniscus
Lomako
Fruth &
Hohmann,
1994
RF
(<500m)
174 - - 5 -
>35
1,155 16.6 16 3-50
Lake
Tumba 107 - - 4-28
Yalosidi GRF 2,380 - - 0-50
Wamba RF 3,353 - 13 0 -
>40
verus community of the Nimba massif (Ivory Coast) build 35.4% of their nests on the ground
(night and day nests were not separated in this study). In Andu (South-West region of
Cameroon), 20% of nest sites of Pan troglodytes ellioti were terrestrial nests (Last & Muh,
2013). In other sites, ground-nesting has rather been described as a rare behaviour (Koops et
al., 2007). The explanation for ground-nesting behaviour may rely in environmental factors
such as low-predation and absence of nesting trees (Last & Muh, 2013), but even more in
cultural or social factors (Koops, 2007). In Babanki-Finge, available nesting trees are
widespread throughout the forest. However, the apparent absence of predators (there is no
leopards in the forest, and chimpanzees are not hunted by humans) would allow chimpanzees
to also nest on the ground. The exclusive arboreal nesting behaviour of Pan troglodytes ellioti
of Babanki-Finge can either be explained by a relatively recent stop of the chimpanzee hunt,
or simply a consequence of a social factor.
69
2.3. Cluster Sizes
No nests were found during the construction processes. Therefore, all nests were discovered,
at the earliest, in the morning after being used by a chimpanzee. Consequently, the size of
clusters had to be estimated. Nest group counting can be biased in several ways. First it is
almost impossible to differentiate two nests built one day apart. Secondly, the distance
between nests estimated to be in a single sleeping site is variable: Fruth & Hohmann (1994)
estimate “close nests in the same stage of decomposition” should be considered as forming
one cluster, while Tutin et al. (1997) argue arguing the nests should be less than 50m apart,
and Dupain et al. (2004) estimate nesting groups are composed of nests within 20m from each
other. The assessment of cluster sizes can therefore be tricky, especially in high density areas
such as Fintse where 21 nests were found within less than 5ha, but where one cannot see
further than 20m due to the rugged landscape and the dense vegetation.
Unlike gorilla groups in which all individuals of the same family can sleep a few meters apart
(Matthews & Matthews, 2004), chimpanzees tend to sleep in very small clusters (Fruth &
Hohmann, 1994), independently on the size of the fusion-fission groups (which have been
censed to up to 100 individuals: Rowe, 1996). The sizes of nest groups between different Pan
communities are compared in Table 15.
Fruth & Hohmann (1994) have pointed out the differences of nesting site sizes between Pan
troglodytes (preferring sleeping alone or in groups inferior to 5) and Pan paniscus (preferring
sleeping in larger groups, median=7 nests per sleeping site). He added the example of the
community of Lopé (Gabon) where 53% of chimpanzees sleep alone, against the group of
bonobos of Lomako (DRC) where 96% of the individuals were observed to nest in groups of
2 or more (generally between 2 and 13 nests per sleeping site). However, the nest groups of
Pan troglodytes ellioti reported by Sommer et al. (2004) (mean=5.7; sd=0.3), of Pan
troglodytes schweifurthii recorded by Ogawa et al. (2007) (mean=5.4) and of Pan paniscus
observed by Reinartz et al. (2008) in Yongo (DRC) (mean=2.3) tend to suggest these
differences may not be significant.
In comparison with other sites, the chimpanzees of Babanki-Finge (mean=1.39) seem to sleep
in group sizes comparable to the community of Pan troglodyes troglodytes from a zone of
Campo-Ma‟an (Cameroon) with a high degree of anthropomorphic pressure (mean=1.30)
(Matthews & Matthews, 2004). In a different part of the Campo-Ma‟an forests with a low
degree of anthropomorphic pressure, the individuals of a community of central chimpanzees
70
Table 15: Comparison of the sizes of sleeping sites between the nest groups of Babanki-
Finge and the ones of Pan trogodytes and Pan paniscus from other sites. The habitat
characteristic codes are the followings: SW: Savannah-Woodland; RF: RainForest;
GWF: mosaic of Grassland and Woodland Forest.
Subspecies COUNTRY Study Site Author Habitat
(Altitude)
Number
of Nests
Nests per Group
Mean Median Range
Pan
troglodytes
verus
Senegal - Baldwin et
al., 1981 SW 83 - 4 1-18
Pan
troglodytes
ellioti
Nigeria Gashaka-Gumti Sommer et
al., 2004
SW
(300m-
2400m
- 5.7 5 1-23
Cameroon
Takamanda
Sunderland-
Groves et
al., 2003
RF
(0m-
1500m)
- 2.86 2.5 -
Bechati-Andu Last &
Muh, 2013
RF
(200m-
1700m)
113 - - 1-14
Bamenda
Highlands
Doumbé,
2013
GWF
(1500m-
2300m)
71 1.39 1 1-4
Pan
troglodytes
troglodytes
Cameroon
Campo-Ma‟an
forests (low
anthropomorphic
pressure) Matthews
&
Matthews,
2004
RF
(<1000m)
137 1.8 -
1-7 Campo-Ma‟an
forests (high
anthropomorphic
pressure)
59 1.3 -
Republic of
Congo Lake Télé
Poulsen &
Clark, 2004 RF 486 2.47 - 1-11
RCA Dzanga-Ndoki Blom et al.,
2001
RF
(340-
615m)
- 2.8 2 -
Equatorial
Guinea -
Baldwin et
al. 1981 RF 66 - 2 1-12
Gabon
Lopé
Fruth &
Hohmann,
1994 RF
(<1000m)
31 - 1->23
Tutin et al.,
1997 424 2.3
Hall et al.,
1998 84 2.95 - -
-
Tutin &
Fernandez,
1984
- 1,606 1.8 1.96 1-10
71
Table 15 (following): Comparison of the sizes of sleeping sites between the nest groups of
Babanki-Finge and the ones of Pan trogodytes and Pan paniscus from other sites. The
habitat characteristic codes are the followings: RF: RainForest; GWF: mosaic of
Grassland and Woodland Forest. *: located in the Salonga National Park.
Subspecies COUNTRY Study
Site Author
Habitat
(Altitude)
Number
of Nests
Nests per Group
Mean Median Range
Pan
troglodytes
schweifurthii
DRC
Kahuzi-
Biega
Basabose
&
Yamagiwa,
2002
RF/Primary
Forest
(2050-
2350m)
104 4.3 - -
RF/Secondary
Forest
(2050-
2350m)
94 4.3 - -
Hall et al.,
1998
RF
(600m-
3308m)
61 2.0 - -
Kasese 19 1.9 - -
Tanzania Ugalla Ogawa et
al., 2007
GWF
(980m-
1712m)
563 5.4 - 1-23
Pan
paniscus DRC
Lomako
Fruth &
Hohmann,
1994
RF
(<500m)
127 - 7 1->23
Lokofa*
Reinartz et
al., 2008
171 8.0 - -
Etate* 58 6.3 - -
Beminyo* 22 11.0 - -
Isanga* 5 1.3 - -
Yongo* 11 2.3 - -
Ikolo* 3 3.0 - -
where observed to sleep in larger groups (mean=1.8) (Matthews & Matthews, 2004).
Therefore the size of sleeping groups might be related to anthropomorphic pressure. Reinartz
et al (2008) established a different conclusion regarding the variability of the mean nest
groups found in Pan paniscus. According to their results, the average cluster size of Pan
paniscus significantly increased with increasing proportion of Marantaceae forests (the nest-
forest type for bonobos). The mean group size of Pan paniscus would therefore depend on the
type of vegetation of the forest. Pan troglodytes target tree species in which they build nests
(Sanz et al., 2007), thus they might also make clusters depending of the density of these trees.
Almost all the nests of nesting sites were built in separate trees. Only two sleeping sites were
found with nests sharing the same tree: three nests and two nests, in Mendong I. According to
72
the literature, most chimpanzees sleep in individual trees (Baldwin et al., 1981; Fruth &
Hohmann, 1994).
2.4. Tree Species
Notwithstanding its botanical limitations, this thesis supports the previous studies stating Pan
troglodytes preferably builds nests in specific trees (Fruth & Hohmann, 1994; Furuichi &
Hashimoto, 2004; Sanz et al., 2007). Carapa grandiflora was found to be the most used
nesting tree by the chimpanzees of Babanki-Finge (65% of the trees). This species is also the
most preferred by the chimpanzees of the Kalinzu forest (Uganda), representing 30% of all
nesting trees (Hashimoto, 1995). Carapa is a gender of trees recognized as quite important for
chimpanzee populations. In Bossou (Republic of Guinea), young leaves of Carapa sp. have
been reported as being used as drinking tools (Sugiyama, 1995). While Carapa procera nuts
were not recorded as eaten by the Pan troglodytes verus of Bossou, they were described as
commonly consumed in the Nimba mountains of Ivory Coast (Matsuzawa & Yamakoshi,
1996). The seed dispersers of the high trees of this genus are forest elephants (Loxodonta
Africana cyclotis) and pouched rats (Cricetomys sp.) (Nyiramana et al., 2011; Theuerkauf et
al., 2009).
The other recognized taxon was Strombosia sp. Hashimoto (1995) has noted a Strombosia
tree (species Strombosia scheffleri) as the 11th
species the most used by the chimpanzees of
Kalinzu, out of 30. The consumed fruits of the chimpanzees of Babanki-Finge is unknown,
but in general, Pan sp. do not build sleeping nests on fruiting trees, but rather when the trees
are not offering ripe fruits, or on a neighbouring plant (Fruth & Hohmann, 1994).
Unlike bonobos, chimpanzees rarely build their nests in two trees or more. In Gombe
(Tanzania), chimpanzees rarely use materials from more than one tree, and in Lopé (Gabon),
more than 90% of the nests were built on single trees, the remaining being built in two trees,
whereas in Yalosidi ,Wamba and Lomako (DRC), bonobos commonly incorporate more than
one tree to the construction of each nest (Fruth & Hohmann, 1994). In Babanki-Finge, the
percentage of nests built in more than one trees is 2.81%, all of them using two trees.
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2.5. Nesting Site Choice
The choice of nesting sites in wild chimpanzee populations has largely been studied (Furuichi
& Hashimoto, 2004). This behaviour tends to be changed where Pan troglodytes spp. and
Gorilla sp. live sympatrically, in order to prevent competition. In these regions, small nesting
groups of chimpanzees appear to build nests in trees whose fruits are not eaten by gorillas
(Basabose & Yamagiwa, 2002).
Fruth & Hohmann (1994) have argued Pan sp. generally prefer to build nests in dense
vegetation. With all nests (recorded during my field work until May the 15th
) built under a
canopy cover higher than 51%, and 7 out of 9 nest sites constructed in areas with a canopy
cover superior to 76%, our data support this statement.
The choice of building a nest quite close to farms and/or houses, as found in Abong-Phen, has
also been recorded in the Kahuzi-Biega lowland, (DRC), where zones adjacent to human
settlements showed a significantly higher nest density than in the heart of the forest (Hall et
al., 1998). Contrarily, in their study in Gabon, Tutin and Fernandez (1984) have shown great
apes are absent from areas of human activities. This difference could be seen as an adaptation
to the anthropomorphic pressure in Kahuzi-Biega and the Bamenda Highlands, in comparison
to the relatively undisturbed forest of the Gabonese study site.
3. Other Wildlife
3.1. Monkeys
Despite the monkeys being described as common by villagers of Bambui, Finge and Kedjom-
Keku, very few were seen during the data collection in Babanki-Finge. The reason of this lack
of data was (according to the same villagers) because the monkeys come closer to the
plantations during the months of July and August: during the harvesting of maize. The only
monkey species found were two Cercopithecines: the tantalus monkey (Chlorocebus tantalus
tantalus) and the putty-nosed monkey (Cercopithecus nictitans martini). The latter is a social
forest species, very often observed in associations with other taxa such as grey-cheeked
mangabeys (Lophocebus albigena) and crowned guenons (Cercopithecus pogoniass spp.) in
Central Africa (Rowe, 1996). It is considered as very adaptable to anthropomorphic pressure
74
(Tutin, 1999), including hunting (Linder & Oates, 2011). Tantalus monkeys are typical
savannah primates which are known for colonizing cleared forests and savannized woodlands,
and described as the most adaptable of all guenons (Kavanagh, 1980).
Another species of monkey locally named “red monkey” was often mentioned by villagers
and field assistants. The potential species described as the „red monkey‟ could be the patas
monkey (Erythrocebus patas), but no individuals were recorded in Babanki-Finge. According
to Nakagawa (1999), patas and tantalus monkeys live sympatrically throughout most of their
range in tropical Africa, but patas tend to prefer open areas than gallery forests. Maybe the
„red monkey‟ mentioned is a description of Erythrocebus patas which is timidly expanding its
range to the South.
It was not possible to tell if the term “black monkey” generally used by local villagers was
exclusively used for the putty-nosed monkey, or also for Cercopithecus preussi. Because
Babanki-Finge is located within the distribution of Cercopithecus preussi (IUCN, 2013), and
its environment corresponds to the preferred habitat of the species (Butynski & Koster, 1994),
it might be concluded that the absence of Preuss‟s guenon is a consequence of over-hunting,
although I do not have any solid evidence for this assumption. If Preuss‟s guenons are
occasionally found in Babanki-Finge like it was mentioned by Ingram & Nsom Jam (2007),
they may temporarily come from other parts of the highlands. There was no sign of the small
population of black and white colobus (Colobus guereza) presented in Ingram & Nsom Jam
(2007) as occurring in Babanki-Finge. According to „Pa‟ Moses, a hunter killed two
individuals a few years ago which would have led the rest of the group to leave the forest.
Colobines (like mangabeys and apes) are regarded as more sensitive to hunting pressure than
Cercopithecines (Matthews & Matthews, 2002).
Despite being known as a typical savannah monkey gradually colonizing pockets of forests
(Higham et al., 2009) no evidences of baboons were found in Babanki-Finge. My field
assistant David Achomochi mentioned having seen two baboons in Abong-Phen the previous
year, and a couple of villagers argued Papio anubis sometimes raid their crops during the
harvesting season (July-August). A camera trap placed in another fragmented forest in the
North of Kedjom-Keku (uncovered during this field work) caught several baboons. The
reason of the absence of Papio anubis in Babanki-Finge is unknown but might be due to an
over-hunt to prevent crop-raiding (mentioned by a villager), or simply the example of the
gradual and slow colonization of the environment by baboons.
75
The response of sympatric primates to fragmentation is variable (Onderdonk & Chapman,
2000). Tantalus and putty-nosed monkeys are almost certainly the last remaining monkeys of
Babank-Finge, and their number may be relatively high due to a competitive release. The
presence of these two very adaptable species of monkeys, and the absence of other less
adaptable species, highlights the high degree of anthropomorphic pressure on the forest.
3.2. Mammals and Birds
Large mammals have been extinct in Babanki-Finge forest. Despite its home range including
parts of Babanki-Finge (Ingram & Nsom Jam, 2007), no traces of forest buffaloes (Syncerus
caffer nannus) were found during the data collecion, and no villager has stated seeing this
ungulate for years. A species of medium-sized antelope was observed. According to its size
and colours, it was either a sitatunga (Tragelaphus speki) or a bushbuck (Tragelaphus
scriptus), both taxa being described as highly adaptable (Allsopp, 1978; Owen, 1970). This
antelope seems to be the largest remaining non-primate animal in Babanki-Finge. A non-
exhaustive list of the most common non-primate mammals of Babanki-Finge would include
rock hyraxes (Procavia capensis), cane rats (Thryonomys swinderianus) and Cooper‟s
mountain squirrels (Paraxerus cooperi) - endemic to the Nigeria-Cameroon mountains - , as
well as several species of duikers comprising blue duikers (Cephalophus monticola), red-
flanked duikers (Cephalophus rufilatus) and bay duikers (Cephalophus dorsalis).
The diversity of birds in Babanki-Finge is extremely rich. Closed-canopies are occupied by
typical forested species such as blue turacos (Corythaeola cristata), black-and-white casqued
hornills (Bycanistes subcylindricus), yellow-breasted boubous (Laniarius atroflavus), double-
billed barbets (Lybius bidenatus), grey cuckoo-shrikes (Coracina caesia), raptors and
sunbirds; while speckled mousebirds (Colius striatus), black-crowned waxbills (Estrilda
nonnula), black kites (Milvus migrans), black-winged kites (Elanus caeruleus caeruleus) and
francolins (Francolinus sp.) are found in forest clearings and grasslands. In addition, the
Bannerman‟s turaco (Tauraco bannermani) and the banded wattle-eye (Platysteira laticincta),
two endemic bird species of the Bamenda Highlands are commonly observed (particularly in
Ndong-Efuh, Mendong I and Fintse).
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4. Comparing Babanki-Finge with Other Fragmented Forests
The African tropical forest has been recognized as being on an extension phase. Comparisons
between old aerial photographs and recent satellite pictures have shown large expansions of
up to one million hectare of forest in the Centre region of Cameroon (Weber, 2001). The
phenomenon of afforestation of savannahs is however reduced (and even cancelled in some
areas) due to an increasing anthropomorphic pressure. It has for consequences, the
fragmentation of forest patches, involving the isolation of animal populations. Consequently,
monkeys and apes, like other mammals, have to adapt.
Even though fragmentation can be seen as a threat, and as dangerous for the survival of
wildlife (Bergl, 2006), savannahs and open areas separating forest pockets are also positively
used by some typical forested species. Tutin (1999) found the biomass of Pan troglodytes
troglodytes, Mandrillus sphinx, Cercopithecus cephus, Cercopithecus nictitans nictitans and
several ungulates, was significantly more important in galleries than in closed canopy forests
of Lopé National Park (Gabon).
The situation is slightly different in Babanki-Finge. Unlike in Lopé, the fragmented patches of
forest are not close to any large forested area. In the study site of Tutin (1999), the patches of
forest were isolated from the closed-canopy forest core by no more than 500m of open area.
Moreover, the open areas fragmenting Babanki-Finge are occupied by farmers and herders
quite often, which make it more difficult for mammals (which see humans as a threat) to
move from one patch to another. The pockets of forest in Lopé are more than six times
smaller than Alegafor (the smaller ensemble of Babanki-Finge), but their status of protection
and their closeness to a larger forest make it more likely to harbour mammals.
The case of Babanki-Finge is more similar to the fragmented forests of Hoima and to the
patches neighbouring Kibale National Park (Uganda), respectively described by McLennan
(2008) and Chapman et al. (2003). The fragmented patches bordering Kibale were linked to
the forest of the national park before the clearing and the creation of agricultural lands
isolating them in pockets. Nowadays, these mid-altitude patches (~1750m) are surrounded by
smallholder farms and grazing lands (Chapman et al., 2003). Hoima forest is also similar to
Babanki-Finge. Situated in a populated area (95.4people/km²) the forest of Hoima is disparate
and concentrated in valleys along watercourses (McLennan, 2008). As in the fragmented
forest of the Bamenda Highlands, the survival of the chimpanzees of Hoima is due to a
powerful hunting taboo (McLennan, 2008).
77
The fragmentation of these forests has undoubtedly increased chimpanzee-human encounters.
However, while villagers of Babanki and Finge mention Pan troglodytes as rare crop raiders
(in opposition to tantalus monkeys considered as pests), McLennan (2008) reported several
conflicts involving chimpanzees threatening farmers in their land. The only recurrent negative
story about chimpanzees in Babanki-Finge was the killing of dogs when they feel threatened
by the canids.
The fragmented forest outside Kibale shows a drastic decreasing of wildlife, only possessing
half of the primates occurring inside the protected area (Chapman et al., 2003). Moreover,
Chapman et al. (2003) have witnessed a dramatic diminution of species and apparent fertility
rate of most monkeys and apes between 1995 and 2000, related to anthropomorphic pressure.
Outlying chimpanzee populations, such as the communities mentioned above, may be
essential for maintaining gene flows between larger groups inhabiting bigger forests
(McLennan, 2008). Nevertheless, the survival of chimpanzees in such narrow areas highly
threatened by clearings is most unlikely in the long-term (Isabirye-Basuta, 2004). Therefore,
an enhanced protection of Pan troglodytes ellioti of Babanki-Finge is critically needed.
5. For a Better Conservation
The decreasing of biodiversity in the Bamenda Highlands is not a recent problem. In Kilum-
Ijim, the regional extinction of forest buffaloes might have happened at least 20 years ago,
while the extermination of forest elephants would have occurred over 100 years ago (Maisels
et al., 2001). Nowadays, despite its quite large undisturbed woodland cover, this montane
forest is not inhabited by any chimpanzees (Chi, 2007). With a small population of Pan
troglodytes ellioti, the Babanki-Finge forest represents a – weak, but living - hope for the
conservation of the last remaining chimpanzees of the Bamenda Highlands.
Although many ancient societies used to forbid the consumption of chimpanzees, the modern
mix of ethnicities concentrated in cities has changed the habits of people toward wildlife in
Africa (Mesange & Honig, 2012). Modern greed (launched by the increasing of trade with
Europeans since at least the XVIIIth
century) has progressively succeeded to African
traditions, provoking a dramatic impoverishment of wildlife (Tah, 2012). Therefore, the
78
survival of the chimpanzees of Babanki-Finge should be seen as a hope for a better
conservation of the forest, and encouraged by the creation of durable local projects.
5.1. Hopes for Babanki-Finge (?)
The unemployment of the villages surrounding Babanki-Finge is a threat to the wildlife of the
forest. About five professional hunters (gun hunters, dog hunters, and spear hunters) are well-
known in the area. However, more and more new hunters are starting to invade the forest.
Although it is widely-known that hunting chimpanzees is not allowed, hunters are not aware
about the prohibition regarding other animals. Throughout its whole range, the Bannerman
turaco is hunted for its feathers which are attached to hats as an honourable and respectful
physical recognition. Nevertheless, the classification of the Cameroonian wildlife law
categorized this bird as an A class animal, and therefore its hunt is clearly prohibited. In
addition, B class animals - such as the bay duiker and Tragelaphus sp. – are carelessly hunted
in the forest, whereas a permit is, by the national law, required for killing any animal of this
category. Other mammals, either classed Least Concerned or Vulnerable (IUCN, 2013), seem
to be overhunted: putty-nosed monkeys, tantalus monkeys and duikers. A control over
hunting is therefore essential to prevent the decline of the wildlife of Babanki-Finge.
Habitat destruction is also a problem. Furthermore, this problem is emphasized in valleys
were the landscape spreads anthropomorphic sounds.
Nevertheless, despites its fragmentation and the anthropomorphic pressure upon it, the
Babanki-Finge forest remains one of the last viable montane forests of Cameroon (Tropek,
pers. comm), and one of the last areas with Pan troglodytes ellioti in the North-West region of
Cameroon (Morgan et al., 2011). Moreover, the superimposition between the satellite pictures
taken by Google Earth©, have shown, despite the high level of anthropomorphic pressure, the
edges of the forest have not really changed for the past three years. The uniqueness of this
forest is armoured by an encouraging local will to protect it. Led by Ernest Vunan, a group of
people from Babanki-Finge is very determined to protect their heritage and to conserve the
wildlife in their forest. A system of tree planting has been created in order to link the patches
of Fintse-Mendong I and Kubuh, which would connect two of the most used patches by the
chimpanzees and obviously other animals. Seeds and young plants are brought from the forest
79
to a nursery in Kedjom-Keku where they grow waiting to be planted back in the mountain.
Hundreds of trees have been planted in the past two years.
The best proof of the effect of this local awareness and will to save the Babanki-Finge forest
and its wildlife, is the fact chimpanzees are still living in it despite its fragmentation. Several
villagers (from Babanki and Finge) have reported encountering in their field a big primate
“walking on its legs like a human”, with “no hair on the face like us” and then fleeing at sight.
Another interesting fact came from a villager from Kedjom-Ketingo (Small Babanki) who
clearly told me it was a taboo for Babanki people to hunt chimpanzees. According to him, it
would be like killing a man. However, he was he only person who mentioned it as a
traditional taboo. Therefore, I cannot conclude if it is either an old traditional taboo (in which
case it would explain the survival of this small population of chimpanzees) or if this „taboo‟
story is relatively recent, coming from the law prohibiting the killing of chimpanzees
(established in 1994: Chi 2007). This second case would be stimulating as it would prove
indigenous populations can assimilate national laws against poaching, and include them to
their traditions.
A better management of the forest and an awareness on which species to hunt are needed in
order to protect the site of Babanki-Finge. To help to fund long-term conservation projects,
tourism can be a useful tool (Macfie & Williamson, 2010). Once mediatised, the Bamenda
Highlands (described as one of the most important regions of endemism in the Cameroon line:
Blackburn et al., 2010) should gain a better recognition as a touristic site.
5.1.1. Touristic Potential
Long-term support for the conservation of the Babanki-Finge forest and its wildlife can be
enhanced by tourism (Macfie & Williamson, 2010). About 10km north from Bamenda, the
Babanki-Finge area is ideally located. This montane forest is situated a few kilometres from
Bambui (via a permanent and well-maintained road) which is placed on one of the most
touristic sites in the English-speaking part of Cameroon: the Ring-Road. This famous road
forms a circle in the North-West region, passing through an important variety of landscapes
and climates (due to the changes of altitude). Different types of tourisms could be created,
attracting tourists for bird watching, site-seeing, and primate tourism.
80
Bird Watching
A rich and diverse birdlife is observable in Babanki-Finge. Internationally recognized as an
important site for birdlife (Ingram & Nsom Jam, 2007; Ndenecho, 2011; Reif et al., 2007) the
Bamenda Highlands, and especially the Babanki-Finge forest, have a lot to win from a better
touristic vision. Villagers from Babanki, Finge and Bambui have much interest in
emphasising the bird richness of their forest, and remaining heritage.
Site-Seeing
Situated in a hilly landscape with high mountains (up to over 2200m), the Babanki-Finge
region presents many natural sites. Water falls (Figures 26 & 27) and caves with large groups
of Egyptian rousettes (Roussetus aegyptiacus) (Figure 28) and Noack‟s roundleaf bats
(Hipposideros ruber) (Figure 29) are widespread all over the forest patches and grasslands.
Figure 26: Photograph of a water fall in Fintse. Credit: Osiris Doumbé.
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Figure 27: Photograph of a water fall in Mendong I. Credit: Osiris Doumbé.
Figure 28: Photograph of an Egyptian rousette caught in Ndong-Efuh, held by one of the
field assistants David. Credits: Arthur Sniegon.
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Figure 29: Photograph of a Noack‟s roundleaf bat caught in a cave in Ndong-Efuh.
Credits: Arthur Sniegon
Primate Tourism
This type of tourism would be the most difficult due to the extreme sensitivity of the primates
of Babanki-Finge. It would need to be preceded by a relative habituation of chimpanzees by
local monitors, and by a controlled hunting over monkeys. These two factors remain
challenging as the habituation of apes is known for being quite difficult and controversial
(Williamson & Feistner, 2003), and a control over monkey hunting would go against deep
traditional habits. Moreover, primate tourism, and particularly great ape tourism, has to be
well-structured (Macfie & Williamson, 2010). It is essential to observe a health protocol to
prevent the transmission of pandemic human viruses to chimpanzees (Homsy, 1999). Careless
behaviours could regrettably provoke a decline in the populations of apes in the forest
(Köndgen et al., 2008). Lastly, the habituation of Pan troglodytes ellioti in Babanki-Finge
must be carefully evaluated as such experiments in areas with a high level of human density
have very often shown increasing rates of chimpanzee-human conflicts (McLennan & Hill,
2010).
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5.2. Conservation in the North-West Region of Cameroon
It is crucial to include traditional rulers and consider traditions in any project of wildlife and
environment protection, to result in efficient and accepted conservation. The western vision of
conservation brought by European and American NGOs has failed, and it is now essential to
look for a different approach, closer to African societies and respected rulers (Ofir Drori pers.
comm.). The awareness of several local chiefs has already brought changes, and particularly
in the North-West region of Cameroon. Fon Doh Ganyonga III, traditional ruler of Bali-
Nyunogha, aware of the necessity to protect wild animals, has banned an old tradition which
consisted in rewarding hunters for the killing of rare or endangered species (Mfonfu, 2006).
Evidence of the increasing concern for the protection of wildlife has also been observed in the
village of Nsem, in the same region. The traditional ruler of the village, Fon Mbonifor, has
handed over a captured olive baboon to the wildlife officials of the North West Regional
Delegation of Forestry and Wildlife as a contribution to conservation effort (Ofir Drori pers.
comm.).
Despite these improvements and increasing awareness, the conservation of wildlife in the
North-West region of Cameroon is facing several challenges. Firstly, the region includes
some of the most populated areas of tropical Africa (Tropek & Konvicka., 2010). Secondly,
there are only two protected areas and none of them has been elevated into a national park: the
Northern part of the Kagwene Gorilla Sanctuary, and the Kimbi Wildlife Reserve (Figure 30).
Moreover, past over-hunting has impoverished the wildlife and wiped out the populations of
large mammals of the latter (Maisels et al., 2001). The protection and management of non-
protected areas is therefore urgently needed.
The forest pockets of Hoima (Uganda) are situated between the Budongo and Bugoma forests
which are both classified as protected areas. It was proposed to maintain the mosaic of
savannah-forest of Hoima, creating corridors in order to permit the gene flow between the
stable populations of chimpanzees of Budongo and Bugoma (McLennan, 2008). A similar
scheme could be made in the Bamenda Highlands. The populations of Pan troglodytes ellioti
of the North-West region of Cameroon are far less known, less stable and less protected than
the populations of Pan troglodytes schweinfurthii of North-West Uganda, and the linking
between forested areas is essential for the survival of this chimpanzee. The forest of Kilum-
Ijim (Figure 30) does not harbour any chimpanzees anymore (Wilkie et al., 2011), but
possesses the largest montane forest of West Africa (over 8, 500ha: Forboseh et al., 2003) and
84
a rich and diverse biodiversity, with several endemic taxa (Cheek & Csiba, 2000). Therefore,
it is a high priority site for the conservation of plant (Maisels et al., 2000) and mammal
diversity (Verheyen et al., 1997). Kom-Wum Forest Reserve (Figure 30) is a protected area
located on the North-West of Babanki-Finge. This forest would possess the highest
chimpanzee density of the North-West region of Cameroon (Morgan et al., 2011). A
replanting tree project roughly linking Kilum-Ijim and Kom-Wum via Babanki-Finge would
be an essential first step for a better conservation of the wilderness in this part of the country.
The last chimpanzee of Kilum-Ijim was killed in the late 1980s (Maisels et al., 2001) and
little is known about the situation and distribution of Pan troglodytes ellioti in Kom-Wum
Forest Reserve (Morgan et al., 2011). Therefore, awareness campaigns need to be conducted
in these areas before any corridor project. Moreover, surveys are needed to be conducted in
order to have a better idea of the status and distribution of Pan troglodytes ellioti and other
Haplorhines in these two protected areas.
The next step of this ambitious project for the survival of one of the richest wildlife of
Cameroon would be to create a multi-site protected area throughout the Bamenda Highlands,
as proposed by Morgan et al. (2011). This project would include and link the forest reserves
of Kom-Wum, Fungom and Ako-Mbembe (Figure 30). It has been proposed because of the
community engagements, and the expected viability of conservation and comprehensive
management in Kom-Wum and Ako-Mbembe. Nevertheless, a lot of work is needed to be
done before reaching this objective. Recently, a preliminary survey on apes has reported a
very low density of chimpanzee nests and monkeys in Fungom Forest Reserve (Ekinde et al.,
2005). Areas must be a minimum secured from poaching before any linking between them, in
order to prevent any expansion of animal trafficking.
The positions of Bafut Ngemba and „Lac Benakuma‟ (Figure 30) are essential and important
for the creation of a multi-site protected area. However, little is known about these two forest
reserves.
85
Figure 30: Map of the mentioned areas of the North-West region of Cameroon. In light
green are shown the forest reserves, in dark green the wildlife reserves, and in orange,
the wildlife sanctuaries. The forests of Babanki-Finge and Kilum-Ijim are also
represented in green polygons. In the middle of the map is visible the „Ring Road‟. This
map is the zoomed and modified map of the Cameroon‟s Forest Estate from the World
Resources Institute: http://www.wri.org/project/interactive-forest-atlas-cameroon.
6. Future Research
In order to manage and protect the environment of Babanki-Finge efficiently, more research is
needed.
6.1. Babanki-Finge
The borders of Babanki-Finge are difficult to establish, mostly because of its fragmentation
and the fact that several villages share the forest. In this thesis, the borders used were the ones
Kilum
Ijim
Kom
Wum Kagwene
Babanki
Finge
Lake Benakuma
Bafut
Ngemba
Kimbi
Fungom
Ako
Mbembe
86
recently delimitated during the informal monitoring of chimpanzees sponsored by John
DeMarco. Nevertheless, because the monitors come from Kedjom-Keku and Finge, the
borders of the forest were limited to these two villages, even though forested patches seem to
still occur on the other side of the mountain (Figure 31). The creation of maps of some
potential other fragments in the North-East of Babanki-Finge is therefore needed to
understand better the spread of the forested patches in the area. Chimpanzee populations
living in fragmented forests have a very large home range, using open areas to move between
them (Onderdonk & Chapman, 2000). Consequently, surveys on chimpanzees and monkeys
are also needed in this side of the mountain, including in the Mbi crater (Figure 31).
Figure 31: Satellite photograph of the East of Babanki-Finge. Kubuh Mendong I, Fintse,
and Mendong II are encircled in orange. The red polygons show some potential
unexplored neighbouring patches. The blue polygon encircles the Mbi crater. Picture
taken from the software Google Earth©.
Surveys on the Haplorhines of Babanki-Finge need to be done during the rainy season and for
a period higher than three months in order to have a better idea of the populations of monkeys
and apes all year long.
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6.2. The North-West Region of Cameroon
Surveys are needed in the forest of Kilum-Ijim, and in the Kom-Wum, Ako-Mbembe,
Fungom, Lac Benakuma and Bafut Ngemba forest reserves in order to do the inventory of the
diversity of monkeys and the survival of the remaining populations of primates in the North-
West region of Cameroon. Wherever ape populations are found, follow-ups will be needed,
and studies on the ecology and survival of the communities of haplorhines will be essential to
establish the dynamic of the species to plan a better protection. Finally, a large campaign of
awareness and sensitivity towards the forest and the animals, as well as a heavy replantation
of trees will have to be emphasized.
6.3. Recommendations
In conclusion, I recommend the following steps for an improved protection of Pan troglodytes
ellioti and its environment in the North-West region of Cameroon:
A longer survey for the diurnal primates of Babanki-Finge is needed to cover both
rainy and dry seasons and to do a complete inventory of the species of monkeys
inhabiting the fragmented forest.
A sensitization campaign in the villages of Kedjom-Keku, Kedjom-Ketingo, Finge and
Bambui about the protection of the forest, and regarding the species of animals which
are prohibited to kill or which require a permit for hunting.
The linkage of the patches of Babanki-Finge via the abandoned land covered with
fern.
The survey of the remaining pockets of forest surrounding Babanki-Finge is urgently
needed in order to establish the distribution of the small population of Pan troglodytes
ellioti in this small area.
Inventories of the mammals inhabiting the Ako-Mbembe, Fungom, Kom-Wum, Lac
Benakuma and Bafut Ngemba reserve forests, the montane forest of Kilum-Ijim, and
the Kimbi wildlife reserve are essential to complete the map of the mammalian
wildlife of the North-West region of Cameroon.
A large sensitization campaign in communities neighbouring the forest reserves of the
North-West region about the protection of the forest, and regarding the species of
animals which are prohibited to kill or which require a permit for hunting.
88
The creation of corridors linking the Babanki-Finge and Kilum-Ijim forests, the Kimbi
wildlife reserve, the Kagwene Gorilla Sanctuary and the forest reserves of the North-
West region of Cameroon.
The achievement of the cited points would be jointly done by an improvement of the
local touristic infrastructure.
7. General Conclusion
The high level of anthropomorphic pressure has almost wiped out all the montane forests of
West-Cameroon, a critical massif for the African biodiversity. This thesis contains the first
maps of Babanki-Fing, one of the last remaining montane forests of the Bamenda Highlands.
Composed of three main ensembles of patches, this forest is fragmented by grasslands and
farms (mostly maize, banana trees and djama-djama). All these pockets of forest are
characterized by being concentrated along streams and rivers. Nevertheless, they vary in sizes,
levels of degradation, and distributions of animals. The observed animals are mostly very
adaptable species, signs of a high level of anthropomorphic pressure. The most Southern
ensemble (Alegafor) – which is the most disturbed - is apparently only inhabited by birds and
duikers, while the most Northern set of patches (including Fintse, Kubuh and Mendong I and
II) is also occupied by Tragelaphus sp., rock hyraxes, putty-nosed monkeys, tantalus
monkeys, and chimpanzees.
The Nigeria-Cameroon chimpanzee is the most endangered taxon of Pan sp. The distribution
of the subspecies is highly fragmented throughout its whole range, and almost unknown in the
North-West of Cameroon. This thesis highlighted the nesting areas (and parts of the nesting
ecology) of Pan troglodytes ellioti of Babanki-Finge. In this forest, chimpanzees seem to
prefer the patches of Fintse and Mendong II as sleeping sites. Most nest groups were found to
be below three nests on average, confirming the fact chimpanzees tend to sleep in very small
clusters, independently on the size of the group. The low height of the nests in trees (below
10m on average) was supposed to come from the high altitude. Because of the shortness of the
field work and the highly varied rate of decomposition of chimpanzee nests, the methodology
failed in assessing the total population of great apes of Babanki-Finge. The remaining
89
chimpanzees of the forest are expected to be very few (less than 20), living in a population not
viable in the long-term.
Further surveys are needed to complete this first inventory of the Haplorhines of Babanki-
Finge. In addition, the conservation of the forest is urgently needed to prevent the
deforestation and hunting rates. Because of a good location on the touristic Ring Road, if
mediatised, the touristic potential of Babanki-Finge could attract tourists for its birdlife and
beautiful landscape. This income should then be wisely used to enhance the development of
the villages, and to improve the protection of the forest.
A project of the use of this mosaic of grassland/forest as a corridor for Pan troglodytes ellioti
to reach larger forests such as Kilum-Ijim and Kom-Wum remains the fragile hope for the
survival of the last chimpanzees of Babanki-Finge.
This thesis contributes significantly to the protection of Pan troglodytes ellioti and of its
natural environment. Hopefully, this first study on a little studied forest and a neglected
population of chimpanzees will attract the international scientific interest in the faith of the
rarest Pan troglodytes in one of the last montane forests of West Africa.
90
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