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THE SPATIAL AND SOCIAL ORGANIZATION OF
FOREST BUFFALO (SYNCERUS CAFFER NANUS)
AT LOPE NATIONAL PARK, GABON

By

LISA MARIE KORTE

A DISSERTATION

Submitted to
Michigan State University
in partial fulﬁllment of the requirements
for the degree of

DOCTOR OF PHILOSPHY

Department of Zoology

2007

ABSTRACT

THE SPATIAL AND SOCIAL ORGANIZATION OF
FOREST BUFFALO (SYNCERUS CAFFER NANUS)
AT LOPE NATIONAL PARK, GABON

By
LISA MARIE KORTE

The Congo Basin forests of central Africa are home to one of the least-studied
large mammals, the forest buffalo (Syncems caffer nanus). Though several
comprehensive studies have been completed on the familiar Cape buffalo
(Syncerus caffer caffer), few data exist for forest buffalo. Data are needed for this
forest—dwelling subspecies to aid conservation planning in the Congo Basin. This
region became a global conservation priority in 1999, when six heads of state
from the Central African nations signed the Yaounde Declaration, promising to
conserve 10% of their nations’ forests as protected areas. In 2002, the president
of Gabon fulﬁlled this promise when he created Gabon’s ﬁrst national parks. One
of these new national parks is Lope National Park, where I spent two years
studying forest buffalo. The goal of my research was to provide basic information
on habitat preference and social structure of forest buffalo.

l tracked nine radiocollared adult female forest buffalo between December
2002 and December 2004. Home ranges of these buffalo averaged 4.55 km2 in
area; the percent of home range overlap between individual radiocollared buffalo

was small. Distance analysis of habitat use from radiotracking data was used to

assess forest buffalo habitat selection at two spatial scales. At the landscape
scale, buffalo selected savanna and marsh habitat over forest habitat within a 72-
km2 study area. Thus, forest buffalo home ranges were savanna-dominated
despite the greater amount of forest habitat available at the park. At the scale of
home range (2.30 km2 to 7.64 kmz), habitat selection within home ranges varied
with season. Adult female forest buffalo preferred forest habitat between March
and August but preferred marsh to forest between September and February.
Forest buffalo dwell in forest habitat, feed on savanna grasses, and wallow in
marshes, utilizing all habitat types within the landscape. Although the subspecies
is forest-dwelling, forest buffalo depend on open habitat adjacent to continuous
forest.

Eighteen forest buffalo herds used the study area with an estimated
population of 342 individuals (~ 5 buffalo/kmz). Mean group size for these 18
herds was 12 :t 2 (range of means = 3-24), considerably smaller than Cape
buffalo herds. For eight radiocollared forest buffalo, mean group size was stable,
varying little throughout the day, seasonally, or between savanna and marsh
habitat. However, herd size varied widely across herds, from fewer than ten
individuals in the smallest herds to more than 20 buffalo in the largest. Large
herd size is associated with home ranges that contain substantial areas of open

habitat, and thus more food resources than forested habitats.

Copyright by
Lisa Marie Korte
2007

ACKNOWLEDGMENTS

I am grateful to Barbara Lundrigan, the chair of my committee, for her
encouragement and support during my dissertation. The advice and expertise of
Kay Holekamp, Catherine Lindell, and Scott Winterstein, the other members of
my graduate committee, were invaluable for completing this research. I also wish
to thank the Michigan State University (MSU) Department of Zoology, the African
Studies Center, the Graduate School, International Studies and Programs, the
US Student Fulbright Program, and the Wildlife Conservation Society (WCS) for
ﬁnancial support. I appreciate the technical advice provided by the Center for
Statistical Training and Consulting on statistical analyses and Judy Olson on map
design. The MSU Museum gave me a comfortable and productive place to work.

I would like to thank the government of Gabon for permission to work at
Lope National Park and for the permits to radio-collar buffalo. I am especially
thankful to the Lope National Park staff for their efforts to preserve the ﬂora and
fauna of the park. WCS ﬁeld veterinarians, Billy Karesh and Mike Kock, handled
the sedation of buffalo for radio collar placement with commendable technical
expertise and respect for the buffalo. Jean Thoussaint Dikangadissi, Edmond
Dimoto, Kath Jeffery, and Ruth Starkey enthusiastically assisted Billy and Mike
with the radio-collaring. I am indebted to Beatrice Kiene Boussoughou, my ﬁeld
assistant, for collecting telemetry data.

The Lopé research station, Station d’Etudes des Gorilles et Chimpanzés

(SEGC), provided me with a place to live, ﬁeld assistants, and vehicles, and

facilitated logistics during my ﬁeld work. I especially appreciate the initial
invitation from Lee White and Kate Abernethy to study buffalo at Lopé and their
logistical and technical support in the ﬁeld. I wish to thank my many colleagues at
SEGC for their helpful suggestions and willingness to always lend a hand with
activities. I am thankful to the parent organization of SEGC, Centre Internationale
de Recherche Médicales de Franceville (CIRMF), for their core support of the
research facilities at SEGC and their acceptance of my study within their
research program.

The US Student Fulbright Fellowship Program, the Hensley Scholarship of
the MSU Department of Zoology, and WCS contributed funds toward my
research in Gabon. Special thanks to the staff of these organizations for their
administrative assistance.

Throughout this degree program, my colleagues, friends, and family
provided consistent support for my research and pursuit of a career in
international wildlife conservation. Colleagues at WCS invited me to regional
meetings and gave me an opportunity to share thoughts with a diverse group of
conservation professionals. It has been a privilege for me to be part of the WCS
team. Faculty and students at MSU provided useful comments on my
manuscripts and presentations. The Molloy and Korte families have been a
constant source of love and encouragement. Finally, I would like to thank my
husband, David for joining me in Lansing for the duration of my dissertation. I

dedicate this dissertation to David and thank him for his patience and love.

vi

TABLE OF CONTENTS

LIST OF TABLES ................................................................................................. ix

LIST OF FIGURES ............................................................................................... x

INTRODUCTION .................................................................................................. 1
Overview of chapters ......................................................................................... 3

Chapter |

HABITAT SELECTION AT TWO SPATIAL SCALES AND DIURNAL ACTIVITY

PATTERNS OF ADULT FEMALE FOREST BUFFALO ........................................ 8
Introduction ........................................................................................................ 8
Materials and Methods .................................................................................... 11
Results ............................................................................................................. 20
Discussion ....................................................................................................... 24

Chapter II

VARIATION OF GROUP SIZE AMONG AFRICAN BUFFALO HERDS IN A

FOREST-SAVANNA MOSAIC LANDSCAPE ..................................................... 39
Introduction ...................................................................................................... 39
Materials and Methods .................................................................................... 42
Results ............................................................................................................. 47
Discussion ....................................................................................................... 50

Chapter III

CALVING AND INTER-BIRTH INTERVALS OF FOREST BUFFALO AT LOPE

NATIONAL PARK, GABON ................................................................................ 61
Introduction ...................................................................................................... 61
Materials and Methods .................................................................................... 61
Results ............................................................................................................. 62
Discussion ....................................................................................................... 63

CHAPTER IV

HERD-SWITCHING IN ADULT FEMALE AFRICAN FOREST BUFFALO

SYNCERUS CAFFER NANUS ........................................................................... 66
Introduction ...................................................................................................... 66
Material and Methods ...................................................................................... 66
Results ............................................................................................................. 67
Discussion ....................................................................................................... 67

vii

CHAPTER V
PERSPECTIVES ON THE ROLE OF BUFFALO IN CONSERVATION AT LOPE

NATIONAL PARK, GABON ................................................................................ 70
Introduction ...................................................................................................... 70
Forest buffalo ................................................................................................... 72
Lopé National Park Management Plan ............................................................ 74
Park management, research, and training institutions at Lope National Park..74
History of buffalo research at Lope .................................................................. 75
Lope NP Management Plan Objectives ........................................................... 76
Summary of recommendations ........................................................................ 84

REFERENCES ................................................................................................... 87

viii

LIST OF TABLES

Table 1.1. Home range estimates based on 100% MCP (minimum convex
polygon) and LoCoH (local area nearest-neighbor convex-hull) methods for nine
radio-collared adult female (AF) forest buffalo (Syncerus caffer nanus) at Lopé
National Park, Gabon, during December 2002-December 2004, including number
of locations, k, and habitat composition of home ranges (ranked by home range
size in ascending order) ........................................................................ 36

Table 1.2. Range overlap between individual adult female (AF) forest buffalo
based on LoCoH home range area estimates. Values are the percentage of
the range of a buffalo (row) shared with another buffalo (column) ................... 37

Table 1.3. The ranks and pair-wise habitat comparisons based on (a) mean
distance of a forest buffalo from habitat Almean random distance to habitat A
within the study area, (b) mean distance of a forest buffalo from habitat Almean
random distance to habitat A within the animal’s LoCoH home range area by
season ............................................................................................... 38

Table 2.1. Effect of time of day, season, and habitat type (marsh or savanna), on
size of forest buffalo groups that contained radiocollared adult females (AF) at
Lope National Park, Gabon, December 2002-December 2004 ....................... 57

Table 2.2. Correlation coefﬁcients (r) between mean and maximum size of group
and home range characteristics based on adult female buffalo with a radio collar
(N=8) ................................................................................................. 58

Table 2.3. Comparison of mean group size estimates for telemetry data versus
data collected on circuit surveys for nine radiocollared adult female buffalo at
Lopé National Park Gabon, December 2002-December 2004 ........................ 59

Table 3.1. Time between births for nine radio-collared adult female (AF) forest
buffalo (Syncerus caffer nanus) at Lopé National Park, Gabon, December 2002-
December 2004. ................................................................................. 65

Table 5.1. Summary of objectives in the Lopé National Park Management Plan
(2006-2011) with management recommendations for forest buffalo and the
potential beneﬁts that could result from implementation of the

recommendations ................................................................................. 86

LIST OF FIGURES
Figure 1.1. Geographic location of study area in Lopé National Park, Gabon ..... 32

Figure 1.2. Locations (points), 100% MCP (black polygons), and LoCoH (color
polygons) home range estimates for 9 adult female forest buffalo (AF1-AF9;
Syncerus caffer nanus) with radiocollars at Lopé National Park, Gabon,
December 2002 — December 2004 ......................................................... 33

Figure 1.3. Mean proportion of locations (:I: SE) during daylight hours in each
habitat by season for 7 radiocollared adult female forest buffalo at Lope National
Park, Gabon, December 2002-December 2004. ....................................... 34

Figure 1.4. Mean proportion of time (:I: SE) spent in each activity category by
habitat type (A) Marsh and (B) Savanna, and time of day for 7 radiocollared adult
female forest buffalo at Lope National Park, Gabon, December 2002-December
2004 .................................................................................................. 35

Figure 2.1. Mean group size of forest buffalo herds at Lopé National Park,
Gabon, (September 2002-December 2004) based on circuit survey data. Means
are based on observations of >2 buffalo for 249 circuits. N=the number of
observations, including observations with and without radiocollared buffalo.
Observations include all age classes. Mean herd size =11.91:t1.55. Range of
group size for each herd was: A (3); B (3-4); C (3-5); D (3-7) E (3-8); F (7-8); G
(5-12); H (3-14); I (3-17); J (4-24); K (3-24); L (5-24); M (4-23); N (3-26); 0 (4-
26); P (4-30); Q (3-43); and R (10-44). AF6 was consistently observed with Herd
P until August 2004, when she was ﬁrst observed with Herd C, in which she
remained through the end of the study

peﬁod ................................................................................................ 53

Figure 2.2. Mean group size of forest buffalo sightings that contained a
radiocollared adult female (AF) at Lopé National Park, Gabon, December 2002-
December 2004. N=the number of observations, representing all observations of
radiocollared buffalo including sightings on circuit surveys and telemetry and
opportunistic observations. Observations include all age classes. Group size
range for each buffalo: AF6c (1-8); AF8 (2-12); AF4 (1-18); AF5 (1-25); AF6p (1-
30); AF1 (2-29); AF3 (5-24); and AF2 (6-46) .............................................. 54

Figure 2.3. a) Home range of each radiocollared buffalo in the study area at Lopé
National Park, Gabon (December 2002 — December 2004); b) Maximum size of
group based on observations that included a radiocollared adult female (AF)
forest buffalo in relation to the area of open habitat in the home range of those
females. Each data point represents maximum group size (N=8) deﬁned by the
presence of a radiocollared buffalo. Positive correlation between maximum group
size and areas of open habitat is signiﬁcant (t=6.35, P<0.05). ....................... 55

Figure 2.4. Seasonal distribution of forest buffalo group size at Lopé National
Park, Gabon, based on circuit survey data. The y-axis represents the number of
observations in each group size class divided by the total number of circuits per
season. Mean group size for all sightings of buffalo, including solitary buffalo,
was 1110.34 based on 775 observations during 249 circuits .............................. 56

xi

INTRODUCTION

My dissertation examines the spatial and social organization of forest buffalo
(Syncerus caffer nanus), a subspecies of African buffalo. This forest-dwelling
subspecies of African buffalo differs markedly in geographic range, morphology,
and behavior from the familiar Cape buffalo (Syncerus caffer caffer). The species
range is throughout Sub-Saharan Africa. Forest buffalo occur in the Congo Basin
forests of central Africa whereas Cape buffalo are on the savannas of eastern
and southern Africa. Body size of forest buffalo (250-320 kg) is approximately
half that of the 400-800 kg Cape buffalo (Haltenorth & Diller 1980). Horn shape
and horn orientation of the two subspecies also differ. Forest buffalo have small,
swept-back horns without the lateral extensions of Cape buffalo horns.
Behaviorally, forest buffalo herds are smaller than Cape buffalo herds. Average
herd size in Cape buffalo is 350 individuals, with the largest herds reaching into
the thousands (Mloszewski 1983; Prins 1996; Sinclair 1977). Group size of forest
buffalo has been estimated at 20 based on survey data in the colonial literature
(Jarrnan 1974). Kingdon (1982) suggests that the suite of differences observed
between these two subspecies of African buffalo reﬂects adaptations for living in
savanna versus forest habitat. However, this has not been tested since our
knowledge of forest buffalo is limited in part due to the logistical challenges of
studying forest animals (Blake 2002).

The goal of my research was to provide basic information on habitat
preferences and social structure of forest buffalo. Though several comprehensive

studies on Cape buffalo have been completed throughout its range (Conybeare

1

1980; Funston et al. 1994; Halley & Mari 2004; Halley & Minagawa 2005; Halley
et al. 2002; Hunter 1996; Mloszewski 1983; Prins 1996; Ryan et al. 2006; Sinclair
1977; Taolo 2003), comparable data for forest buffalo are lacking. Information on
forest buffalo comes mostly from surveys of African forest mammals (Blake 2002;
Prins & Reitsma 1989; Sidney 1965; Tutin et al. 1997; White 1994). However,
one recent study successfully tracked a single herd of forest buffalo at Dzanga-
Ndoki National Park in the Central African Republic (Melletti et al. 2007a). Like
Cape buffalo, forest buffalo are grazers (i.e., they feed almost exclusively on
grass) yet the geographic range of the forest buffalo is primarily forest, where
grasses are limited to open areas along rivers or streams, and in forest clearings
or small savanna patches within the forest landscape (Blake 2002; Melletti et al.
2007a). This presents interesting ecological questions about forest buffalo that
can only be addressed by observational research that reveals their habitat use
and social structure.

Another impetus for understanding forest buffalo ecology is to inform
conservation initiatives in the Congo Basin. Despite the lack of information on
forest buffalo behavior and ecology, this subspecies is highlighted as important
for conservation planning in the Congo Basin forests (Blake 2002; Coppolillo et
al. 2004; Melletti et al. 2007a; Melletti et al. 2007b). The creation of new national
parks in the region was inspired by the recognition of the Congo Basin forests as
a global conservation priority (Kamdem-Toham et al. 2003). In Gabon, Lopé
National Park is one of the largest of thirteen national parks created in 2002 to

protect the forest and unique wildlife of the region. At Lopé NP, buffalo are one of

the most abundant large mammals of the forest-savanna mosaic in the northeast
corner of the park (Tutin et al. 1997). Forest buffalo are clearly visible in the
savanna habitat during the September through November wet season (Molloy
1997). This visibility could tie in with tourism to generate revenue for the Lopé NP
if the population is managed effectively. However, few studies have focused on
forest buffalo (Alers & Blom 1988; Molloy 1997) and park managers lack the
basic information needed for managing forest buffalo. The success of the newly
created national parks will depend on park staff acquiring the data needed to

sustain wildlife populations and make sound management decisions.

Overview of chapters
In Chapter I, I investigate how forest buffalo use habitat. It seems unlikely that
forest buffalo, a large grazing species, would have a geographical range that is
heavily forested. Grass, the primary food source of buffalo, is widely scattered in
small ﬂushes in the forest landscape. Kingdon (1982) suggests that small
clearings found within forests could support forest buffalo because the humid
climate provides sufﬁcient food resources for this large grazer throughout the
year. Indeed, forest buffalo concentrate in forest clearings and along rivers and
streams within the forest, and are rarely seen where clearings or water are
absent (Blake 2002; Melletti et al. 2007a). In Chapter I, I examine habitat
selection at two spatial scales for eight radio-collared forest buffalo at Lopé NP.
First, I compare home range area composition to the overall landscape. Then, I
compare locations of buffalo within their home range area to the habitat

composition of the home range. I demonstrate that buffalo feed in open areas,

wallow in marsh habitat, and increase use of forest habitat when food resources
in open areas are low. Although the geographic range of forest buffalo is the
forest habitat of central Africa, forest buffalo depend on open habitat adjacent to
forest habitat for feeding and resting sites. The work presented in Chapter I has
been accepted for publication in the Journal of Mammalogy.

The objectives of Chapter II were to document group size in forest buffalo
at Lopé NP and examine the inﬂuence of season, habitat, and time of day on
group size, factors that are known to inﬂuence group size in other ungulates
(Borkowski & Furubayashi 1998; Brashares & Arcese 2002; Hirth 1977; Jannan
1974; Marchal et al. 1998). Mean herd size of Cape buffalo is 350 individuals
with a range of 12 to more than 1,500 individuals (Prins 1996; Sinclair 1977). In
Chapter II, I report group size of forest buffalo at Lopé NP. Eighteen forest
buffalo herds used the study area with an estimated population of 342 individuals
(~ 5 buffalo/kmz). Mean group size for the 18 herds was 12 1: 2 (range of means
= 3-24), considerably smaller than Cape buffalo herds. For eight radiocollared
forest buffalo, mean group size was stable, varying little with time of day, across
seasons, or between savanna and marsh habitat. However, herd size varied
widely across herds, from fewer than ten individuals in the smallest herds to
more than 20 buffalo in the largest. Large herd size is associated with home
ranges that contain substantial areas of open habitat, and thus more food
resources than forested habitats. At Lopé NP, food resources probably have
more inﬂuence on group size of forest buffalo than predation due to the low

predation pressure on adult animals and enforced hunting restrictions.

In Chapter III, I report on inter-birth intervals and gestation period of forest
buffalo. Since little is known about calving and inter-birth intervals in forest
buffalo, these data are valuable despite the small sample size. Recent work by
Ryan et al. (2007) on reproductive ecology in African buffalo describes a longer
gestation period for Cape buffalo than would be expected given their body size.
They suggest the protracted gestation period is an adaptation to the seasonal
environment. Of the nine radiocollared adult female buffalo, gestation period
could be determined for only one. The ten-month gestation period observed was
slightly shorter than that reported for Cape buffalo (11.5 months; Sinclair 1977). I
found an inter-birth calving interval of two years with 67% calf mortality. The high
mortality rate combined with long inter-birth intervals may contribute to low
reproductive rates, and should be noted by park managers. Calf survival seems
to be inﬂuenced most strongly by the availability of food resources at Lope NP,
but further studies are needed. Chapter III is in press at the African Journal of
Ecology.

In Chapter IV, I examine herd switching among nine radiocollared adult
female forest buffalo. In Cape buffalo, adult females, subadults and juveniles live
in mixed herds throughout the year, whereas adult male Cape buffalo move
among mixed herds or form bachelor herds when not attached to a mixed herd
(Mloszewski 1983; Prins 1996; Sinclair 1977; Turner et al. 2005). Adult female
Cape buffalo were not known to switch herds. However, recent studies of Cape
buffalo in Botswana and South Africa report evidence of herd-switching in adult

female Cape buffalo (Cross et al. 2005; Halley et al. 2002). In Chapter IV, I report

that only one of the nine radiocollared adult female buffalo at Lope NP switched
herds during the two-year study period. Also, forest buffalo had well-deﬁned
home ranges with small areas of overlap. Unlike Cape buffalo, there was little
evidence of herd-switching among adult female forest buffalo, suggesting a more
stable social structure in forest buffalo.

In Chapter V, I present a plan for integration of park management and
capacity building for wildlife managers with long-term ecological research of
forest buffalo. Conservation planning for species will be most successful when an
understanding of an animal’s ecology can be integrated into the management
objectives of the parks and reserves in which it lives. In this ﬁnal chapter, I
examine the relationship between forest buffalo (Syncerus caffer nanus) and the
objectives presented in the Lopé National Park Management Plan 2006-2011.
Forest buffalo, one of the most abundant large mammals on the savannas of
Lope NP, have the potential to be a focus of research, management, and training
activities. Because the results of these activities could increase revenue for the
park and increase funding for conservation activities, I consider how a long-term
conservation study of forest buffalo could beneﬁt each park objectives. An
institutional collaboration between the Lopé research station and the training
center to promote this study could make a signiﬁcant contribution to both park
management and the training of scientists in Gabon. Study results from Lopé NP
could be applied to protected areas throughout the Congo Basin Forest Region

of central Africa.

Chapter |

Korte, L. 2008. Habitat selection at two spatial scales and diurnal activity patterns
of adult female forest buffalo. Journal of Mammalogy 89.

Chapter I

HABITAT SELECTION AT TWO SPATIAL SCALES AND DIURNAL ACTIVITY
PATTERNS OF ADULT FEMALE FOREST BUFFALO

Introduction

Ungulates are the focus of many studies of mammalian ecology and
space use. Antelope are of particular interest due to their considerable variation
in morphology, geographic range, and behavior among the species and the
number of species (i.e., 143 species within Family Bovidae). Because wild
populations are observable and captive populations are easily managed, the
group is well-studied. Early studies highlighted important relationships between
feeding style and habitat use by African antelopes (Gwynne & Bell 1968; Jarman
1974). Subsequently, advanced statistical techniques and a phylogeny of extant
species have been used to verify the signiﬁcant correlation between habitat use
and feeding styles in ungulates (Pérez-Barberia et al. 2001). However, few
studies have examined habitat use for single species that range across a variety
of habitats (Brashares & Arcese 2002; Brashares et al. 2000; Jannan 1974).
Data are lacking for analysis of habitat selection for those species that may
provide the most insight on the relationship between feeding style and habitat
use. Here, I investigate space use of the African buffalo (Syncems caffer). This
large-bodied grazer (Halley & Minagawa 2005; Hofman & Stewart 1972; Janis
1988; Jarrnan 1974) has both savanna and forest—dwelling populations

(Haltenorth & Diller 1980; Kingdon 1997; Sinclair 1977).

Differences in morphology, geographic range, and behavior are evident
among subspecies of Syncerus. The 2 most widely recognized subspecies are
the savanna or Cape buffalo (Syncerus caffer caffer), and the forest buffalo
(Syncerus caffer nanus; Grubb 1972; Wilson & Reeder 2005). The more familiar
Cape buffalo are massive animals, weighing 250-800 kg, with downward curved
horns (Haltenorth & Diller 1980; Sinclair 1977). At weights of 250-320 kg, forest
buffalo are approximately one-half the size of Cape buffalo (Haltenorth & Diller
1980). Forest buffalo have small, swept back horns without the lateral extension

characteristic of Cape buffalo.

Cape buffalo have been studied in most detail in the savannas of eastern
and southern Africa where vast, open grasslands support herds averaging 350
animals, with large groups reaching into the thousands (Conybeare 1980;
Funston et al. 1994; Halley & Mari 2004; Halley & Minagawa 2005; Halley et al.
2002; Hunter 1996; Mloszewski 1983; Prins 1996; Ryan et al. 2006; Sinclair
1977; Taolo 2003). In contrast, the geographic range of S. c. nanus is the Congo
Forest Basin region of central Africa (Haltenorth & Diller 1980; Kingdon 1997;
Sinclair 1977). Due to the forest habitat and their elusive lifestyle, few data exist
for S. c. nanus (Blake 2002; Melletti et al. 2007a). Relative to Cape buffalo, much

less is known about space use of forest buffalo.

Although some studies of forest buffalo have addressed space use,
conclusions are limited due to short study periods and small sample sizes. At
Lope National Park, Gabon, Molloy (1997) found that forest buffalo fed on the

ﬂush of new grass on savannas, but data collection was limited to 1 6-month

9

period and home range estimates relied solely on visual observations of only 3
groups. Though Abernethy (K. A. Abernethy, Station des Etudes des Gorilles et
Chimpanzés, Gabon, in litt.) later tracked radiocollared forest buffalo over a 2-
year period at Lopé NP, the sample size (i.e., 1 adult female and 1 adult male)
was too small for analysis of habitat selection (Aebischer et al. 1993). At Dzanga-
Ndoki National Park, Central African Republic (CAR), forest buffalo were highly
dependent on forest clearings, but observations were of a single group (Melletti
et al. 2007a). For analysis of habitat selection, home range data with larger

sample sizes are needed.

Kingdon (1982) suggested that small open areas found within forests
could support forest buffalo because the humid climate provides sufﬁcient food
resources for this large grazer throughout the year. For mammals, habitat
productivity and body mass inﬂuence areas of home ranges (Harestad & Bunnel
1979; McNab 1963). Home range size increases with body mass (McNab 1963);
however, smaller home ranges are found in more productive habitats (Fisher &
Owens 2000; Harestad & Bunnel 1979). Although ﬁeld surveys indicate that
forest buffalo use open habitat within the forest landscape (Blake 2002; Prins &
Reitsma 1989; Tutin et al. 1997; White 1994), data on space use within home

ranges are needed to test if forest buffalo select open areas.

In this study, I used radiotelemetry to examine diurnal space use of adult
female forest buffalo at Lopé National Park, Gabon. My objectives were to 1)
estimate home range areas, 2) examine ranging patterns, 3) assess habitat use

at 2 spatial scales, and 4) record diurnal activity of forest buffalo. I compare my

10

ﬁndings for forest buffalo with observations of Cape buffalo and analyze how

forest buffalo use habitat within the forest landscape.

Materials and Methods

Study area—l observed buffalo in Lopé National Park, Gabon. This 5000-
km2 park is located in the center of Gabon just south of the equator (S 00° 12' 04;
E 11°36’ 05; Fig. 1.1). White (1983) classiﬁes the park as lowland tropical
rainforest. Average total annual rainfall is 1500 mm, unevenly distributed over 2
dry seasons (December-February and June-August) and 2 wet seasons (March-
May and September-November). Temperatures ﬂuctuate little throughout the
year with maxima ranging from 27-31°C and minima from 20-22° C (White &
Abernethy 1997).

The ungulate community at Lopé NP includes forest buffalo, forest
elephant (Loxodonta cyclotis), sitatunga (Tragelaphus spekir), bushbuck
(Tragelaphus scriptus), red river hog (Potamochoerus porous), water chevrotain
(Hyemoschus aquaticus), and 7 species of duikers (Cephalophus; Tutin et al.
1997). Leopards (Panthera pardus), the only large carnivore at Lope NP, are a
natural predator of the ungulate community. Forest buffalo were 1 of the main
prey items found in leopard scat; however, based on analysis of bones in the
scat, leopards apparently prey only on juvenile buffalo (Henschel et al. 2005).
Hunting pressure by humans on the population of forest buffalo is low because
the park headquarters is adjacent to the study area and because tourists and

researchers use the area on a daily basis.

11

The research site included 3 habitat types: marsh, savanna, and forest.
Digital maps of the northeast area of the park are available to researchers at the
Lopé ﬁeld station, Station d’Etudes des Gorilles et Chimpanzés (SEGC), in the
form of ArcView 3.x (Environmental Systems Research Institute 1999) shape
ﬁles, and include the locations of roads, rivers, marsh, savanna, and forest
habitat types. For marsh habitat that had not been previously mapped, I
measured the perimeter of marsh habitat, created shape ﬁles, and added the
marsh shape ﬁles to the digital map collection at SEGC. Thus, maps including

marsh, savanna, and forest habitat were available for analyses.

Observations of buffalo were restricted to the mosaic of savanna, marsh,
and forest habitat in the northeast corner of the park. The majority of the park is
forest habitat and large mammal surveys by White (1994) suggested there were
few buffalo in the forest adjacent to the mosaic habitat. When present, large
mammals are clearly visible in the savanna habitat. Sixty kilometers of road
traverse this mosaic and are used by tourists and researchers. Park
management staff annually burn the savannas, maintaining the open areas that
would otherwise be colonized by forest (Peyrot et al. 2003). These open areas
facilitate the viewing of large mammals on safari drives, and provide scenic vistas
for tourists. Savanna areas are burned in small sections based on natural ﬁre
breaks and roads in the landscape. Small patches are burned as soon as the
rains have stopped each year, when grasses are sufﬁciently dry to burn well. A
few small areas are burned each week during the 2"“ dry season through the end

of September or beginning of October, depending on rain and burning conditions.

12

This patchy burning creates flushes of new grass, and the number of buffalo
observed in the savanna areas is low in July but increases in August and
remains high through December (based on observations between July and

December 1996; Molloy 1997).

Observations of animals and data sampling.— The ﬁrst 3 months of
observation (September-December 2002) were used to identify groups of forest
buffalo. Three ﬁxed road circuits were established to monitor buffalo, so that
individuals from different groups could be selected for radiocollaring. These
circuits covered all open savanna areas visible from roads and each circuit was
surveyed at least 4 times each month (twice during the morning, 0630-0930 h,
and twice during the late afternoon, 1530-1830 h). Date, time, location, group
size, and activity of individuals were recorded. Age class and sex were
determined for the majority of individuals in each group. If there was a clear view
and buffalo were within 50 m of the observer, a Sony digital video camera (model
DCR-TRV140 NTSC) was used to record buffalo. Videos were reviewed to note
additional unique characteristics that could be used to recognize individuals.
Distinct pelage characteristics and horn morphology were used to differentiate
individuals. Using easily recognizable buffalo as indicator animals, it was

possible to distinguish groups.

Radiocollars were placed on 8 adult female buffalo (AF1-AF8) during the
1“"t week of December 2002. Four collars were manufactured by HABIT, Inc.
(Victoria, British Columbia) and 4 by Telonics, Inc. (Mesa, AZ); receivers from

both companies were used to track buffalo. Females were from 8 different

13

groups, which represented about one-half of the groups using the study area.
Ideally, Borger et al. (2006) recommend that telemetry studies strive to maximize
the number of animals with radiocollars; however, I was limited by the number of
collars. Therefore, I chose to sample animals within the same sex and age class
(i.e., adult females) because core group members in Cape buffalo are adult
females (Mloszewski 1983; Prins 1996; Sinclair 1977). Sampling additional sex
and age classes during this study would have severely reduced sample size

(Aebischer et al. 1993).

Two collars malfunctioned with 1 failure in July 2003 and a 2"cl in October
2003. In early December 2003, the collar on AF3 that failed in October 2003 was
replaced. A 2"“ collar was placed on a new adult female (AF9) in the same group
as the remaining buffalo (AF5) with the malfunctioning collar. The 2"“ collar failed
immediately resulting in 1 group with 2 collared buffalo (AF5 and AF9), but
neither collar emitting a signal. Without the signal, it was not possible to locate
animals when they were in forest habitat. Observations were therefore limited to
visual observations of these 2 animals for the 2"“ year of the study. Hence, data
for adult females AF5 and AF9 were used only for the analysis of home range
overlap and were excluded from the analyses of habitat selection and activity
patterns. I therefore report results for 7 adult females over a 2-year period for the

analyses of habitat selection and diurnal activity patterns.

During 3 study periods (December 2002 through April 2003, July through
December 2003, and July through December 2004), radiocollared animals were

located twice per week on different days. To ensure that animals were tracked

14

throughout the day, daily observations were blocked into four 3-hour
observations periods (i.e., 0630-0930, 0930-1230, 1230-1530, and 1530-1830).
Each animal was tracked at least twice during each of 4 daily observation periods
over the course of a month. However, observations were reduced to 1 location
per week between 0900-1500 during May through June 2003 and January
through June 2004. Opportunistic observations of radiocollared animals outside
the search periods were also recorded. For each observation, date, time,

location, group size, and activity state of individuals were recorded.

Activity data were collected during observations when radiocollared
animals were visible. Focal animal surveys on radiocollared individuals lasted 20
min with an activity recorded every minute. A group scan was conducted if the
collared animal was not alone; the activity of each animal in the group was
recorded every 5 min for 20 min. Observations were only recorded during
daylight hours. Visual observations were also limited to open savanna or marsh
areas because observing animals in the forest resulted in animals either running
from or toward the observer. In both cases, behavior was changed due to the

observer.

All research was conducted under a permit from the Gabon Ministry of
Water and Forests and under the direction of SEGC, the ﬁeld station of the
Centre Internationale de Recherche Médicales de F ranceville (CIRMF). Work
with live animals was carried out in a humane manner and in accordance with the

guidelines of the American Society of Mammalogists (Gannon et al. 2007) and

15

Michigan State University. Veterinarians from the Wildlife Conservation Society’s

Field Veterinary Program handled the buffalo sedation for radiocollar placement.

Determination of home ranges—I plotted the locations of radiocollared
animals as points in an ArcView 3.x (Environmental Systems Research Institute
1999) shape ﬁle. Each collared animal had a point shape ﬁle, with date, time
seen, time of ﬁrst and last telemetry ﬁx, activity, number of buffalo in group,
notes, departure time, and type of observation (i.e., visual or ﬁx) recorded into

the attribute table.

I used the local nearest-neighbor convex-hull construction (LoCoH; Getz
and Wilmers 2004) to estimate the area of the home range for each radiocollared
buffalo, as well as Minimum Convex Polygons (MCP) for comparison with
previous studies. Locations used for the estimations included visual observations
of the animals and telemetry ﬁxes and were statistically independent because
observations were recorded during different time periods if recorded on the same
day (an infrequent occurrence) or different days for the majority of observations. I
used the animal movement analysis ArcView extension (Hooge & Eichenlaub
1997) to determine the MCP and to calculate its area. While the MCP method is
an accepted standard for calculating ranges and is widely used because of its
simplicity (Burgman & Fox 2003; White & Garrott 1990), it has been criticized
because presence of outliers can dramatically overestimate the home range area
(Burgman & Fox 2003). Thus, LoCoH has been advanced by Getz and Wilmers
(2004) and was 1St used to estimate ranges for African buffalo in South Africa

(Ryan et al. 2006).

16

I used the LoCoH (Local Convex Hull) Homerange Generate ArcView
extension to estimate area of the home range for each radiocollared buffalo (see
Getz and Wilmers 2004 and Ryan et al. 2006 for details). This extension uses the
locations to create the convex hull with each location and its k nearest neighbors.
Because the k parameter is user-selected, I ran this method for k values from 2
to 40 to identify the plateau that gives stable-area values across a range of k
values that represent the estimated area of the home range. When several
plateaus occurred, I chose k values that eliminated the unused areas within the
range because the topology of the study area did not include lakes, mountains,
or inhospitable habitats that may be avoided by buffalo. This selection process
followed the “minimum spurious hole covering” rule (Getz & Wilmers 2004) and I
report k values for estimated LoCoH home range areas. I used a Mann-Whitney-

U test to compare the estimated areas of the 2 methods (LoCoH and MCP).

I also examined home range overlap between individuals and between
years for each individual. To determine whether radiocollared buffalo maintained
discrete home ranges, I calculated the percentage of home range overlap
between radiocollared buffalo based on areas calculated using the LoCoH
method. Home range overlap between years for individual radiocollared buffalo
was examined with the Mann-Whitney-U test. This test also was used to test for

differences in the area of home ranges between years.

Habitat measurements and landscape variables. —To measure available
habitat in the overall landscape, a 72-km2 study area was delineated based on

the merged LoCoH home range areas of the radiocollared buffalo with a 1-km

17

buffer. The habitat composition of the study area was 2.45% marsh, 43.20 %
forest, and 54.35% savanna. Forest habitat within the study area included forest
fragments, which have been described by Tutin et al. (1997): 1) gallery forest
(narrow strips of forest along watercourses continuing at one end to join the main
forest), 2) “corridor” (a narrow gallery linked at both ends to forest), and 3)
bosquets (small forest blocks completely surrounded by savanna; Fig.1.2).

Forest habitat in the study area also included continuous forest.

Habitat selection and distance analysis. —Several methods exist to
compare resource use and availability (Thomas & Taylor 2006); I used Euclidean
distances to assess nonrandom habitat use and to rank the habitat types
(Conner & Plowman 2001). The distance analysis is robust to telemetry error and
unlike compositional analysis can include zero-use areas without inﬂuencing
Type I error (Bingham & Brennan 2004). Each radiocollared buffalo was treated

as 1 sample.

I used the Nearest Features ArcView extension (v. 3.8a; Jenness 2004)
to calculate distances between animal locations and the nearest representative
of each habitat type; distances for each animal were averaged for the analysis. I
examined both 2nd order selection of habitat within the landscape and 3rd order
selection of habitat within the home range (Johnson 1980). For comparison at the
landscape level, 2,500 random points were generated in the study area and
distances between each habitat type and each random location were calculated
and averaged to create an average distance for each habitat type. I calculated

ratios for each animal by dividing the distance associated with buffalo locations

18

by distances derived from random locations within the study area. I used a
MANOVA to determine if the mean ratio vectors differed from a vector of 1, which
indicates non-random use. To test for signiﬁcant differences between individual
ratios for each habitat type and the available habitat in the study area, univariate
t-tests were used. Habitats were ranked using pair-wise habitat comparisons to

construct a ranking matrix of t-statistics.

For analysis within the home range, locations of buffalo were paired with a
random location within each home range and distances to each habitat for the
random locations were calculated. For each season, the ratios for each animal
were calculated by dividing the buffalo locations by distances derived from
random locations within the home range. Then I used the distance analysis as
described above to determine if use was signiﬁcantly different from random
locations and to rank habitats. All statistics for the distance analysis were
performed in SAS (version 9.1; SAS Institute Inc. 2002) using code provided by

Conner and Plowman (2001) with q=0.05.

Diumal activity pattem.—l used MANOVA in SAS (version 9.1; SAS
Institute Inc. 2002) to examine diurnal activity patterns for the 7 adult female
forest buffalo with functioning radiocollars, testing for overall effects of habitat,
year, season, and time of day on activity (feeding, active, and inactive). I tested
for signiﬁcant differences attributable to habitat and time of day among the
means for each of the 3 behavioral categories. I used analysis of variance (proc

GLM in SAS) followed by LSD (least square difference) to test for signiﬁcant

19

differences in activity categories of buffalo between savanna and marsh habitat

type and among time periods.

Results
Ranging pattems.—The MCP and LoCoH methods for calculating home
range areas yielded similar results with no signiﬁcant difference between the
estimated home range areas for the 2 methods (MCP N=7, LoCoH N=7, U=29,
P=0.31). However, because the MCP method is more likely to overestimate
home range areas (e.g., for buffalo AF4 and AF6, Fig. 1.2), LoCoH area
estimates are reported and used for analyses. Please note that Figure 1.2 in this

dissertation is presented in color.

Radiocollared adult female forest buffalo maintained home ranges that
were less than 8 km2 and of the same size and in the same location from year to
year. Mean home range area was 4.55:0.72 kmz, with individual areas ranging
from 2.30 km2 to 7.64 km2 for the 7 collared buffalo included in the analysis
(Table 1.1; Fig. 1.2). The total number of locations was not correlated with
estimated home range area (12:0.23, P>0.05, N=7), indicating that increased
sample size would not have resulted in larger home ranges. Individual home
range areas were not signiﬁcantly different between year 1 (December 2002—
November 2003) and year 2 (December 2003-November 2004; Mann-Whitney
U- test; year 1 N=7, year 2 N=7, U=26, P=0.45) with the percent of home range

overlap between years ranging from 50% to 91% for the 7 individuals.

20

Overfap in home ranges between radiocollared buffalo. —The percent of
home range overlap between individual radiocollared buffalo was small (Table
1.2). The 1 exception was 99% overlap between AF5 and AF9, which were
members of the same group. Four buffalo (AF 1, AF2, AF4, and AF7) had ranges
that overlapped with AF6. She used a long and narrow range, running north and
south in the center of the study area (Fig. 1.2). Thus, the perimeter of her range
had the greatest opportunity to overlap with others, and the highest percentages
of overlap (28%, 18%, and 17%) involved her home range. Excluding AF5, AF6,
and AF9, the percent of home range overlap between any 2 individuals was s
7%, including adjacent ranges where overlap was most often zero or < 1%. The
number of locations in areas of overlap was small and there were few occasions
when 2 radiocollared buffalo were present in the overlapping area at the same
time. Overlap occurred primarily in savanna areas with almost no home range

overlap in forest habitat, especially forest galleries.

Habitat selection based on distance analysis. —The average distance
between the location of a buffalo and the nearest marsh, savanna, and forest
habitat was 141.45 m, 24.49 m, and 66.32 m, respectively. In the study area, the
average distance between randomly selected locations and the nearest marsh,
savanna, and forest habitat was 390.98 m, 115.73 m, and 56.15 m, respectively.
At the landscape level, the analysis of distance ratios indicated that locations of
forest buffalo differed from random locations (F=268.82, d.f. = 3, 4, P<0.0001).
Buffalo were found closer to marsh (t=-13.31, d.f. = 6, P<0.0001) and to savanna

(t=-27.09, d.f. = 6, P<0.0001) than expected based on the relative availability of

21

these habitats in the study area. There was no difference between locations of
buffalo and random points with regard to distance to forest (t=0.87, d.f. = 6,
k0.4179). A ranking of habitats indicated that buffalo were found signiﬁcantly
closer to savanna than marsh habitat and signiﬁcantly closer to marsh habitat

than to forest habitat within the study area (Table 1.3).

The proportion of locations within each habitat type varied with season
(Fig. 1.3); therefore, habitat selection within home ranges was examined by
season. The analysis of distance ratios indicated that locations of buffalo differed
from random locations for all seasons: Mar-May (F=68.49, d.f. = 3, 10,
P<0.0001), Jun-Aug (F=5.89, d.f. = 3, 11, P=0.0120), Sep-Nov (F=7.96, d.f. = 3,
11, P=0.0042), and Dec-Feb (F=5.89, d.f. = 3, 11, P=0.0120). During Mar-May,
buffalo were found closer to forest (t=-7.66, d.f. = 12, P<0.0001) than expected
and associated less with savanna (ff-4.13, d.f. = 12, P=0.0014) than expected;
there was no difference between locations of buffalo and random points with
regard to marsh (t=1.85, d.f. = 12, P=0.0890). Buffalo were found closer to marsh
than expected during Sep-Nov (t=-4.53, d.f. = 13, P=0.0006) and Dec-Feb (t=-
4.12, d.f. = 13, P=0.0012), but associated less with marsh habitat than expected
during Jun-Aug (t=-4.12, d.f. = 13, P=0.0012). During June-February, there were
no differences between locations of buffalo and random points with regard to
distance to savanna (Jun-Aug t=0.92, d.f. = 13, P=0.3755; Sep-Nov F067, d.f.
= 13, P=0.5168; Dec-Feb #092, d.f. = 13, P=0.3755) or to forest (Jun-Aug
#193, d.f. = 13, P=0.0761; Sep-Nov F190, d.f. = 13, P=0.0805; Dec-Feb

t=1.93, d.f. = 13, P=0.0761).

22

Within home ranges, a ranking of habitats indicated that forest was
proportionally used most during March through August when buffalo were found
signiﬁcantly closer to forest than marsh or savanna (Table 1.3). During
September through February, buffalo were located signiﬁcantly closer to marsh
than forest whereas there was no signiﬁcant difference between marsh and

savanna or between forest and savanna.

Diumal activity patterns and behavior—Buffalo tended to use savannas
for feeding and marsh areas for resting during daylight hours (Fig. 1.4). The
mean proportion of time spent feeding varied with habitat (F=29.55, d.f. = 1, 441,
P<0.0001) and time of day (F=5.57, d.f. = 3, 441, P=0.0009); buffalo most often
fed on savannas in the early morning. During daylight hours, buffalo spent more
than 30% of their time feeding. Active behavior also varied with habitat (F=9.03,
d.f. = 1,441, P=0.0028) and time of day (F=4.48, d.f. = 3, 441, P=0.0041).
Buffalo were most active in savanna habitat and in the late morning (0930-1230).
Buffalo spent less than 15% of daylight hours in active behaviors. As would be
expected, periods of inactivity also varied with habitat (F=46.30, d.f. = 1, 441,
P<0.0001) and time of day (F=9.99, d.f. = 3, 441, P<0.0001). Inactive behaviors
were most often observed in marshes in the late afternoon. During daylight

hours, buffalo spent more than 38% of their time inactive.

Proportions of time spent in different behaviors did not differ between
years (F=0.85, d.f. = 6, 874, A=0.99, P=0.5309) or among seasons (F=0.34, d.f.
= 9, 1061.3, A=0.99, P=0.9604). However, behavior varied signiﬁcantly among

individuals (F=2.02, d.f. = 18, 1225.2, A=0.92, P=0.0070). This variation could be

23

attributed to 1 individual (AF 8) that spent a larger proportion of daylight hours
feeding (56%) than inactive (38%). The other 6 buffalo spent a larger proportion
of daylight hours inactive (54%) than feeding (37%). When AF8 was removed

from the model, there was no overall effect of individuals.

While it was not possible to visually monitor animals when they were
tracked to forest areas, it was possible to monitor the signal emitted from the
collar. Over the ﬁrst 18 months of the study, animals were monitored for 20
minutes after pinpointing their location in a forest area. During this time, it was
possible to determine if animals remained stationary or moved away. Based on
303 observations, buffalo remained in place during the 20-min observation period
and were most likely resting in 1 location. Buffalo were relatively inactive during
daylight hours between March and August, when more than 60% of their

locations were in forest habitat (Fig. 1.3).

Discussion
Home range size—Home range size for adult female forest buffalo at

Lopé NP (2.30-7.64 km2) was considerably smaller than home range sizes for
Cape buffalo, which have been reported at 10.50 to 296.3 km2 in eastern Africa
(Prins 1996; Sinclair 1977), and over 1000 km2 in southern Africa (Hunter 1996).
Small home range size relative to that of Cape buffalo was expected based on
the much smaller body size of forest buffalo and the higher productivity of their
habitat, 2 factors that can inﬂuence home range size (Harestad & Bunnel 1979;

McNab 1963).

24

Larger home ranges typically occur in less productive habitats (Harestad &
Bunnel 1979). While habitat productivity is usually expressed in terms of mean
annual net primary productivity or NPP (Murphy 1975), a more available
measure, annual rainfall, was used here, following Harestad and Bunnell (1979)
and Fisher and Owens (2000). Based on available NPP data, regions with high
annual rainfall typically have higher NPP (Murphy 1975 1977). In eastern Africa,
mean home range size of Cape buffalo is < 200 km2 (mean annual rainfall > 600
mm; Prins 1996; Sinclair 1977); however, home range size notably increases to >
1000 km2 (a 5-fold increase) in the arid region along the border of Botswana and
Zimbabwe (mean annual rainfall ~ 500 mm; Hunter 1996). Where NPP data are
available, tropical forest has a higher NPP than savanna (Murphy 1977; 1975)
indicating the potential for patches of grass in a forest opening to provide buffalo

a constant food source.

The forest buffalo at Lopé NP, an area with high annual mean rainfall
(1500 mm), inhabit a much more productive area than do most Cape buffalo
(Conybeare 1980; Hunter 1996; Prins 1996; Sinclair 1977). In addition, the
annual burning of savanna at Lopé NP creates open grass areas that provide
food for buffalo. Home range size of forest buffalo in my study was similar to that
of 2 radiocollared forest buffalo examined in a pilot study at Lopé between
December 1998 and June 2000; in that case, home range size for a solitary adult
male was 4.94 km2 (N=82 locations) and 3.70 km2 for an adult female (N=88
locations; K.A. Abernethy, in Iitt.). Despite the small size of these areas, the

humid climate provides sufﬁcient food resources for this large grazer throughout

25

the year. Thus, home range sizes observed in Lope NP support Kingdon’s (1982)
hypothesis that small open areas found within rain forest could support forest
buffalo, and follow the predicted relationship of home range size to body size and

habitat productivity.

Ranging pattems.—My data demonstrate that groups of forest buffalo at
Lopé NP occupy separate home ranges. Adjacent home ranges showed very
little overlap (typically < 1%) and radiocollared buffalo from different groups were
rarely present in the overlapping area at the same time. Large areas of overlap
were in savanna areas, but buffalo tended to use forest galleries within their
home ranges separately. One individual (AF6) that overlapped considerably in
space with 4 other radiocollared buffalo spent little time in the areas of home
range overlap and moved to the southern portion of her home range via forest
galleries. In contrast, the percent of overlap between the other 4 pairs with
adjacent home ranges was small. Despite distinct home ranges and occasional
encounters, territorial defense was not observed among forest buffalo at Lopé
NP. Similar discrete home ranges with little evidence of territorial defense have
been described for Cape buffalo (Prins 1996; Sinclair 1977). The ﬁnding of
exclusive forest buffalo home ranges at Lopé is consistent with the hypothesis

that a single group of forest buffalo occupies a forest clearing (Blake 2002).

Patterns of home range use by radiocollared buffalo suggest that forest
buffalo are more sedentary than Cape buffalo, for which seasonal movement has
been observed (Funston et al. 1994; Halley & Mari 2004; Halley et al. 2002; Ryan

et al. 2006; Taolo 2003). Adult female forest buffalo were consistently and

26

reliably located in the same areas throughout the 2-year study period whereas
Cape buffalo can migrate 10-40 km between seasons in Botswana (Halley et al.
2002). Cape buffalo return to the same dry-season home ranges (Halley and
Mari 2004) indicating site ﬁdelity, which I also observed for forest buffalo. The
home range of 1 forest buffalo (AF 1) in my study included 96% of the home
range of a single radiocollared adult female buffalo tracked at Lopé between
1998 and 2000 (K.A. Abernethy, in Iitt.). This same area also was used by a
group that I studied in 1996 (Molloy 1997), suggesting that occupation of an area
by groups of buffalo probably extends over longer periods than the 2-year
timespan of my study. Home ranges of forest buffalo in Lopé NP appeared

remarkably stable in both time and space.

Habitat selection at the landscape level. ——My data suggest that forest
buffalo select proportionally more open habitat (marsh and savanna) and use
continuous forest less than would be expected based on available habitat at the
landscape level (Fig. 1.2, Table 1.3). Surveys of large mammals in forests (Blake
2002; Prins & Reitsma 1989; White 1994) have found that forest buffalo are
absent or present at low densities in continuous forest, which similarly suggests
that buffalo avoid continuous forest. For example, in the continuous forest area
adjacent to my study area, mammal surveys over 2 14-month periods covering 2
annual cycles reported few observations of forest buffalo and low estimates of
biomass (White 1994). During March and April, 2 graduate students repeated 1
of the White (1994) transects and found that dung of forest buffalo was absent

from the continuous forest (Bankert & Schenk 2003). Low estimates of biomass

27

or few signs of forest buffalo in continuous forest have been reported from other
forest sites in Gabon and the Republic of Congo (Blake 2002; Prins & Reitsma
1989). My landscape-level results and work on forest buffalo in C.A.R. (Melletti et
al. 2007a) suggest that forest buffalo select open habitat for their home ranges,

where their main food source, grass, is plentiful, rather than continuous forest.

Habitat selection within home ranges—Forest buffalo had stable,
savanna-dominated home ranges, but habitat use within home ranges varied
with season (Fig. 1.3), and for at least a portion of the year buffalo preferred
forest habitat during daylight hours (Table 1.3). Between March and August
>60% of locations of forest buffalo were in forest habitats; however, forest habitat
represented <50% of the area of home ranges of forest buffalo (Table 1.1).
Continual 12-hour monitoring during daylight hours of 3 of the radiocollared forest
buffalo conﬁrmed high use of forest between February and May (Bankert &
Schenk 2003). In addition, buffalo dung was commonly found in forest fragments
in April and May 2003; signiﬁcantly less dung was present in savanna, marsh,
and continuous forest during these months (Bankert & Schenk 2003). Tourists
visiting the park between March and August sometimes leave with an impression
that the park has no buffalo because buffalo primarily use forest habitat during

the daylight hours.

Forest fragments, galleries, and corridors are used by forest buffalo more
often than continuous forest and most individuals based their home ranges in a
single forest fragment or gallery (i.e., 10-33% of the locations for AF1, AF2, AF3,

AF4 and AF6). Adjacent home ranges of radiocollared buffalo had minimal

28

overlap in savanna areas and no, or smaller areas of overlap, in forest fragments
or galleries. In addition, buffalo remained close to the savanna areas and did not
penetrate deep into the forest. Even animals with home ranges on the edge of
the continuous forest remained near the forest edge. This is consistent with the
low biomass of buffalo reported for continuous forest (White 1994) and the high

biomass of buffalo found in forest fragments at Lopé (Tutin et al. 1997).

The March-through-August period of high forest use includes wet and dry
periods with similar temperature ranges, which suggests that forest buffalo use
forest habitat for reasons unrelated to rainfall or temperature alone. Small
streams run throughout the landscape at Lope NP, which eliminates the need for
migration of forest buffalo during the dry season based on limited water
resources. Similarly, Cape buffalo in the Serengeti show little movement within
home ranges, remain near watercourses, and maintain smaller home ranges
than those of sympatric migrating ungulates during the dry season (Sinclair
1977). In areas dominated by forest or bush, Cape buffalo feed exclusively on
grasses and forbs and stable isotope analysis has conﬁrmed that they are not
consuming the leaves of trees or bushes (Halley & Minagawa 2005). Buffalo at
Lope NP moved into the savannas to feed on the ﬂush of new grass when the
savannas were burned at the end of the June-through-August dry season and
were highly visible on the savanna areas between September and February.
Although they dwell in forest habitat, forest buffalo depend on open habitat for
food.

29

Marshes also were an important open habitat between September and
February. Home ranges of forest buffalo were 4% marsh and 19% of locations of
buffalo were in marsh habitat, despite the small percentage of marsh habitat in
the study area (only 2.45% marsh). The mud wallows within marsh habitat
contributed to the preference for marsh habitat, especially between September
and February. After early morning feeding bouts in the savanna areas, buffalo at
Lopé NP often retire to a wallow and rest in the marsh until sunset. Groups
tended to frequent 1 or 2 wallows within their range and it appeared that
repeated use of the same wallow enhanced water and mud conditions. On the
lshasha Plateau of Virunga National Park, Democratic Republic of Congo,
African buffalo use wallows for therrnoregulation and drinking (Mugangu et al.
1995) and, among temperate bovids, bison (Bison bison) wallow to gain relief

from biting insects (McMillan et al. 2000).

Because my analyses were limited to data from diurnal observations of
adult female buffalo, some differences may be observed for male buffalo and/or
nocturnal observations. Males may differ in home range size based on
observations of Cape buffalo, where adult males wander more widely (Halley &
Mari 2004; Prins 1996). However, the 4.94-km2 home range area estimated for
the solitary adult male monitored in a pilot study (K.A. Abernethy, in litt.) fell
within the range of areas for radiocollared adult females reported in my study. In
addition, adult females with radiocollars were in groups that included an adult
male, which was regularly with the group (Chapter II). For the majority of visual

observations, the adult male was with the group, and I had no observations of the

30

identiﬁed males from these groups in locations outside the home range areas of

the groups. For a deﬁnitive result, it would be necessary to track adult males.

Nocturnal locations of radiocollared forest buffalo would most likely not
signiﬁcantly change the location of the home ranges or estimated home range
area. Given the stability of the ranges over the 2 years of the study, it seems
unlikely that animals would leave their ranges only at night, resulting in a location
or size change. But, nocturnal observations may inﬂuence activity patterns and
habitat selection within home ranges. For the majority of Cape buffalo, grazing is
done during the nighttime hours (Prins 1996; Sinclair 1977; Taolo 2003), which
suggests that the early morning peak of feeding observed at Lopé NP may be a
continuation of nocturnal feeding. lf forest buffalo at Lope NP spend more time
feeding at night, then use of savanna habitat would increase whereas use of
forest and marsh would decrease. Therefore, nocturnal observations may
change conclusions about habitat preferences within home ranges, especially if
forest buffalo are using the savanna areas at night during the peak of diurnal use
of forest, i.e., March through August. Regardless of how the overall patterns of
habitat use may change with nocturnal data for activity patterns, the diurnal
observations of adult female buffalo demonstrate that habitat preference changes

with season based on food availability.

31

 

 

    

30 300 3,000
_ Kilometers _ Kilometers _ Kilometers
Lopé National Park Gabon Africa

 

Figure 1.1. Geographic location of study area in Lopé National Park, Gabon

32

 

 

 

 

86 87 88 89 790 91 92 93 94

 

 

 

 

 

 

‘ v ' / " i
as. .31.. _ - 51...“, “is.
\ 5 l I 0 e I
85 "I?! — —~»- Ni: 85
\O ' . o '
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84 7““ ; o v ‘ 84
. ,1- o
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17 J .l "‘ j _
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3 . ' o
._ . .j' ‘7.
79 ' 7 ‘ _, . 79
78 9 - 78
77 I o— 77
1‘ .
76 - 76

86 87 as 89 790 91 92 93 94
AF1 AF2 AF3 ‘AF4 AF6 AF7 AF9
- Marsh D Forest Savanna

Projection: UTM Zone 32
Source: SEGC maps Gn'd labels: 1000-meter UTM 32 S L. Korte 10437

 

Figure 1.2. Locations (points), 100% MCP (black polygons), and LoCoH (color
polygons) home range estimates for 9 adult female forest buffalo (AF1-AF9;
Syncerus caffer nanus) with radiocollars at Lopé National Park, Gabon,
December 2002 - December 2004.

33

 

 

up
u.-

'1"

\p‘.

[w- I ”I*
I
d

1......

m'lu
n
.
.
t
.. 4|
‘ c

 

 

0.8 - DMarsh lSavanna lForest

    

 

 

0.6 -

0.4 -

0.2 ‘

Mean proportion of locations

0.0 ‘

    
 

   

Mar-May Sep-N ov Dec-Feb

wet wet

 

 

  

dry

Figure 1.3. Mean proportion of locations (:1: SE) during daylight hours in each
habitat by season for 7 radiocollared adult female forest buffalo at Lopé National
Park, Gabon, December 2002-December 2004.

34

A) Marsh

 

   
  

 

 

 

0‘8 _ D early am
{Elate am

0.6 _ Iearly pm
I late pm

 

Mean proportion of time

 

 

 

 

 

 

 

 

Feeding Active Inactive
B) Savanna

0.8 1 El ear1y am
E El late am
‘5 l earl m
a 0.6 y p
r: I late pm
.9
t
O
Q.
9
Q.
C
(0
(D
E

 

 

 

 

Feeding Active Inactive

Figure 1.4. Mean proportion of time (:I: SE) spent in each activity category by
habitat type (A) Marsh and (B) Savanna, and time of day for 7 radiocollared adult
female forest buffalo at Lopé National Park, Gabon, December 2002-December
2004.

35

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36

Table 1.2. Range overlap between individual adult female (AF) forest buffalo
based on LoCoH home range area estimates. Values are the percentage of the
range of a buffalo (row) shared with another buffalo (column).

 

 

AF1 AF2 AF3 AF4 AF5 AF6 AF 7 AF8 AF9
AF1 -- 7 O 0 0 26 0 <1 0
AF2 5 --- <1 0 0 <1 0 O 0
AF3 0 1 --- 0 0 0 0 0 0
AF4 0 0 O --- <1 1 6 0 0 0
AF5 O 0 0 <1 --- O 0 0 56
AF6 16 <1 0 7 0 --- 1 0 0 O
AF7 O 0 0 0 0 1 7 --- 0 O
AF8 <1 0 0 0 0 0 0 -- 0
AF 9 0 O 0 0 99 0 0 0 --

 

37

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.00.00..0> 0003.00 0000.0...0 .000...00.0 0 00.0000 A2082.

 

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38

Chapter II

VARIATION OF GROUP SIZE AMONG AFRICAN BUFFALO HERDS IN A
FOREST-SAVANNA MOSAIC LANDSCAPE

Introduction
Comparative studies of group size in African antelope have contributed much to
our understanding of social organization (Jarman, 1974; Brashares, Garland 8.
Arcese, 2000; Shultz & Dunbar, 2006). However, considerable variation in group
size occurs in some species, potentially confounding the interpretation of
interspecies comparisons (Brashares et al. 2000; Jarman 1974). Fortunately, this
intraspeciﬁc variation is, in itself, valuable for elucidating the factors that inﬂuence
group size and structure. Indeed, because populations of a species have a
shared evolutionary history, the inﬂuence of various ecological factors (eg.
resource distribution, predation pressure, and habitat structure) on group size
can be examined without the confounding inﬂuence of phylogenetic effects
(Brashares & Arcese 2002).

African buffalo (Syncerus caffer) are an example of a species that occurs
in habitats ranging from closed forest to open savanna habitat. Cape buffalo
(Syncerus caffer caffer) and forest buffalo (Syncerus caffer nanus) are the two
most widely recognized of the ﬁve Syncerus subspecies (Wilson & Reeder 2005).
Cape buffalo, weighing 400 to 800 kg, are larger than forest buffalo, occur
throughout the savanna areas of eastern and southern Africa (Haltenorth & Diller
1980; Sinclair 1977), and have a mean group size of 350 (range 12 to more than
1,500; Prins 1996; Sinclair 1977). Cape buffalo have been the focus of several

comprehensive studies (Mloszewski 1983; Prins 1996; Sinclair 1977); only a few

39

studies have been done on forest buffalo and the results of those studies are
limited by small sample sizes (Abernethy 1998, 1999, 2000; Melletti et al. 2007a;
Melletti et al. 2007b) and short study periods (5 6 months; Molloy, 1997; Alers
and Blom, 1988). The smaller, forest-dwelling S. caffer nanus (250-320 kg) is
found in the forests of central Africa (Haltenorth 8. Diller 1980; Sinclair 1977).

Few data are available for group size of forest buffalo. Comparative
studies of African antelope use an estimated group size of 20 for forest buffalo
(Brashares et al. 2000; Jarman 1974; Shultz & Dunbar 2006), though forest
buffalo also are reported to occur in groups of eight to ten buffalo (Mloszewski
1983; Sidney 1965). These estimates are based on single observations from the
colonial literature, but one recent study has tracked group size of a forest buffalo
herd during a two-year period. This herd at Dzanga-Ndoki National Park in the
Central African Republic increased from 16 to 24 due to eight births with neither
immigration nor emigration (Melletti et al. 2007b). In the Dzanga-Ndoki herd,
smaller group size was observed during the wet season and the herd frequently
split into two subgroups.

Season, habitat, and time of day are factors known to inﬂuence group size
in ungulates (Borkowski & Furubayashi 1998; Brashares 8. Arcese 2002; Hirth
1977; Jarman 1974; Marchal et al. 1998). Seasonal variation in group size has
been observed in Japanese sika deer (Cervus nippon) with largest group size
during seasons when food resources were optimal (Borkowski 8. Furubayashi
1998). Group size in sika deer increased during the morning and evening

periods, but the authors suggest that these are large feeding aggregations rather

40

than groups with close links (Borkowski & Furubayashi 1998). Similar
aggregations are formed in the European roe deer (Capreolus capreolus) when
deer feed in large open ﬁelds during the morning and evening periods (Marchal
et al. 1998). In fallow deer (Dama dama), the largest groups are formed in open
habitat (Apollonio et al. 1998). Hirth (1977) reported an inverse relationship
between group size and vegetation cover for white-tailed deer (Odocoileus
virginianus).

African buffalo are large grazers, feeding primarily on grass (Halley &
Minagawa 2005; Hofman & Stewart 1972; Janis 1988). On a continental scale,
group size of African buffalo is small in forests relative to savannas (Jarman
1974; Sinclair 1977). This probably reﬂects the distinct differences in quantity and
dispersion of food between these habitats (Jarman 1974). If the quantity and
dispersion of food inﬂuence group size, then group size should vary with habitat
type and across seasons at a local level as well.

Here, I inquired whether season, habitat, or time of day inﬂuence group
size of forest buffalo at Lopé National Park, Gabon. My study area was a forest-
savanna mosaic where the amount and condition of grass varies with habitat and
season (White & Abernethy 1997). Habitat types are distinct in regard to food
resources for buffalo at Lope NP, where forest habitat lacks grass for these large
grazers and grass is abundant in open habitat. The quantity of food increases
during wet seasons, when the savannas have a ﬂush of new grass. I tested three
hypotheses, speciﬁcally asking whether group size is larger (1) during wet

seasons, (2) in savanna habitat, or (3) during early mornings. Group size was

41

expected to increase during wet seasons because of the ﬂush of new grass. If
large groups were feeding aggregations, large group size was expected during
early morning periods when buffalo are feeding in savanna habitats (Chapter I). I
also investigated the relationship between group size and three home range
variables (1) total area, (2) area of open habitat, and (3) percent of the total area
that was open habitat. Herds with the largest home range areas and greatest
amount of open habitat within their home range were expected to have large

group size.

Materials and Methods

Study area—Lope National Park is primarily forest habitat, but the northeast
corner of the park (0° 10' S, 11° 35’ E) is a mosaic of savanna, marsh, and forest
fragments, including gallery forest along streams (T utin et al. 1997; Fig. 1.1). The
72 km2 study area included open (2.45% marsh and 54.35% savanna) and
closed (43.20% forest) habitat. Mean annual rainfall in the study area is 1500
mm, and rainfall varies seasonally. March-May and September-November are
the two seasons with the highest rainfall; December—February is relatively dry,
but the driest period is June-August. Temperatures ﬂuctuate little throughout the
year, with monthly maxima ranging from 26-32° C and minima from 20-22° C
(White 8. Abernethy 1997).

Sources of group size data: radiocollared buﬁ'alo.—In December 2002,
radio collars were placed on eight buffalo (AF1-AF8), representing eight different
herds. Collars were placed on adult females to maximize sample size within one

age and sex class. I selected adult females because adult females form the core

42

of Cape buffalo herds (Mloszewski 1983; Prins 1996; Sinclair 1977) and
preliminary study suggested that social structure of forest buffalo was similar to
that of Cape buffalo (Molloy 1997). Each buffalo was in a group of >3 at the time
of immobilization and collaring. Due to the failure of the collar on AF5, a ninth
collar was placed in December 2003. This ninth female (AF9) was in the same
group as AF5. HABIT, Inc. (Victoria, British Columbia) manufactured ﬁve of the
collars and Telonics, Inc. (Mesa, AZ) manufactured the other four collars; I used
receivers from both companies to track buffalo.

During three periods of the study (December 2002 through April 2003,
July through December 2003, and July through December 2004), radiocollared
buffalo were located twice weekly. To ensure that radiocollared buffalo were
tracked throughout the day, daily observations were blocked into four 3-hour
observation periods (i.e., 0630-0930, 0930-1230, 1230-1530, and 1530-1830).
Over the course of a month, each radiocollared buffalo was tracked at least twice
during each of these four daily observation periods (i.e., a minimum of 8
observations per month). Observations were reduced to one location per week
between 900-1500 during May and June 2003, and January through June 2004
(i.e., a minimum of 4 observations per month).

Group size (i.e., number of individuals including the radiocollared buffalo),
date, time, and location, were recorded for each sighting (i.e., observation) of a
radiocollared buffalo. Buffalo that were within meters of each other and were
coordinated in their activity at the time of observation were considered a single

group (Leuthold 1976). A count of all buffalo per sighting was feasible in savanna

43

and marsh habitat, where buffalo were clearly visible. It was not always possible
to see each buffalo well enough for identiﬁcation to sex and age class, but when
possible age class and sex were determined. For example, when buffalo where
lying in a wallow, it was possible to count buffalo, but not necessarily sex and
age all individuals, especially juveniles and calves. Age classes reported for
Cape buffalo were used to estimate age class of forest buffalo (i.e., calvesz<12
months, juveniles: between 12 and 24 months, subadults: 2-5 years, and adults:
>5 years; Pienaar 1969; Sinclair 1977). Visual observations were limited to
savanna and marsh habitat because observing buffalo in the forest resulted in
animals either running away from or toward the observer, which made it
impossible to obtain an accurate count.

Research was conducted under permit from the Gabon Ministry of Water
and Forests, and under the direction of Station d’Etudes des Gorilles et
Chimpanzés (SEGC), the ﬁeld station of the Centre International de Recherches
Médicales de Franceville (CIRMF). Veterinarians from the Wildlife Conservation
Society’s Field Veterinary Program handled the buffalo sedation for radio collar
placement. Radio collars were left on buffalo for future research.

Statistical analysis: radiocollared buffalo—Analyses were performed
using SAS statistical software (SAS Institute Inc. 2002), and where appropriate,
mean values :I: S.E. are reported. Group size data based on sightings of
radiocollared forest buffalo were analyzed for the effect of individual, season,
habitat, and time of day using a mixed linear model (PROC MIXED). Because

individual had a signiﬁcant effect on group size, the mixed linear model was also

used for group size data of each radiocollared buffalo to analyze the effect of
season, habitat, and time of day on group size. For seasons, the least-squared
means (using LSMEANS within PROC MIXED) were computed and compared to
test for signiﬁcant differences in mean group size between seasons and among
years (eg. Jun-Aug 2003 compared with Sep-Nov 2003, Jun-Aug 2003
compared with Jun-Aug 2004). For pairs of seasons with a signiﬁcant difference,
I used a paired Student’s t—test (oc= 0.05) to compare the mean number of
radiocollared buffalo for which I observed signiﬁcant differences in group size
between seasons to the mean for those with no change in group size between
seasons.

Sources of group size data: circuit surveys—Three ﬁxed road circuits of
23 km, 22 km, and 18 km in length were established to record group size of
buffalo herds without a radiocollared buffalo. Circuits covered savanna and
marsh habitat visible from roads, and each circuit was surveyed at least four
times every month (twice during the morning, 0630-0930, and twice during the
late afternoon, 1530-1830) from September 2002 to May 2003, July to December
2003, and July to December 2004.

Group size was deﬁned as the number of individual buffalo counted per
sighting. Buffalo within approximately 200 meters of one another were treated as
one sighting or observation (i.e., as members of the same group). Sightings of
buffalo were usually separated by more than 1 km; thus buffalo in different
groups were rarely in visual contact with each other. The number of sightings per

circuit ranged from 0-14 (mean=3). For each sighting, group size, date, and

45

location were recorded. When feasible, sex and age class were recorded for
each buffalo as well. Distinct characteristics of individual buffalo (e.g. pelage and
horn morphology) made it possible to distinguish among different social groups
or herds. Buffalo that were repeatedly observed in a single sighting were
identiﬁed as members of the same herd.

Results (mean, maximum, and minimum size of group) from the two
observation methods, (1) telemetry, and (2) circuit surveys for sightings of
radiocollared buffalo were compared using Mann-Whitney-U tests.

Home range characteristics based on radiocollared buffalo—Digital maps
of the northeast area of the park are available to researchers at SEGC in the
form of ArcView 3.x shape ﬁles (Environmental Systems Research Institute
1999), and include the location of roads, rivers, marsh, savanna, and forest
habitat types. I plotted locations of radiocollared buffalo as points in an ArcView
shape ﬁle and used these points with the Local Convex Hull (LoCoH) Homerange
Generate ArcView extension to estimate home range area for each radiocollared
buffalo (Getz & Wilmers 2004; Ryan et al. 2006). I also measured the amount of
each habitat type within each home range area.

Group size variation among radiocollared buffalo was examined in relation
to three home range variables: (1) total home range area, (2) area of open
habitat in the home range, and (3) percent of the total area that was open habitat
in the home range. I used a Pearson’s correlation coefﬁcient (oc= 0.05) to test
whether mean group size for radiocollared buffalo was signiﬁcantly correlated

with each of the three home range variables. Data from two radiocollared buffalo

46

(AF5 and AF9) were excluded from the home range analysis because their
collars failed within six months, and without signals from collars, it was not
possible to track these buffalo to forest locations. The relationships between
maximum group size and each of the three home range variables were also

examined.

Results

Does group size vary?—For radiocollared buffalo, mean group size for all
observations was 1413 (1-46). Mean group size was stable with little variation in
group size between marsh and savanna habitats or throughout the day, but there
was evidence of some variation across seasons. When data from circuit survey,
telemetry, and opportunistic observations of radiocollared buffalo were combined,
signiﬁcant effects on group size of radiocollared buffalo (F[8,374]=99.75;
P<0.0001), time of day (F[3,374]=2.62; P=0.0515), and season (Fp'874]=16.25;
P<0.0001) were observed in the mixed linear model. Only the effect of habitat
(marsh or savanna) was not signiﬁcant (F[.,374]=0.04; P=0.8445). In addition to
the overall model, analysis was completed for each radiocollared buffalo because
of the signiﬁcant differences in group size among the radiocollared buffalo (Table
2.1). No signiﬁcant effect of habitat type on group size was observed for the
radiocollared buffalo; mean group size was 15 for both marsh (1510.48) and
savanna habitat (1510.33). Also, no signiﬁcant effect of time of day on group size
was observed for eight of the nine radiocollared buffalo.

Season had a signiﬁcant effect on group size for ﬁve of the nine

radiocollared buffalo (Table 2.1). Differences between pairs of seasons were

47

consistent among radiocollared buffalo with a tendency for larger group size
during wet seasons than during dry seasons. But, group size was larger during
dry than wet seasons in a few cases. For pairs of seasons showing a signiﬁcant
difference in group size, the mean number of radiocollared buffalo exhibiting no
change in group size (i.e. stable; mean=5.44) was signiﬁcantly greater than the
mean number of buffalo exhibiting a seasonal change in group size (mean=2.56;
t17=-5.88, P=<0.0001).

Variation of group size among herds—Eighteen herds used the study
area with an estimated population of 342 individuals, ~ 5 buffalo/km2 (Fig. 2.1).
The estimated total population is based on the sum of maximum group size for
the 18 herds. All herd members were not together at all times, and observations
of subgroups were common. For herds of fewer than 10 individuals, all buffalo
were known; for larger herds (i.e., >10 animals), the adult males and the majority
of adult females were known. Individuals were repeatedly observed with the
same herds based on these known individuals. For example, all members of
Herd G were identiﬁed and individuals in this herd were only observed with other
buffalo from Herd G. Though buffalo were often observed in subgroups, herd size
was stable. For example, group size for AF8 (Herd G) was consistently small,
and never exceeded 12 during 144 observations (Fig. 2.2). In contrast, AF2 was
consistently observed in a large group (mean=2711; range 6-46; Herd Q). She
was observed with fewer than 10 buffalo during only eight of 78 observations,

and the majority of sightings (65%) included >20 buffalo.

48

Home range characteristics—Because group size did not differ between
marsh and savanna habitat, these two habitat types were combined (i.e. open
habitat) for analysis of home range areas. Both mean and maximum group size
were signiﬁcantly positively related to home range area and to the amount of
open habitat in the home range of radiocollared buffalo (Table 2.2; Figure 2.3).
Percent of open habitat in the home range was not signiﬁcantly related to mean
or maximum group size (Table 2.2).

Method of data collection—Since radiocollared buffalo were observed
during circuit surveys, it was possible to use results from the radiocollared buffalo
to compare telemetry observations with circuit survey observations. Minimum
group size was signiﬁcantly lower for observations made using telemetry than for
groups assessed solely on the basis of circuit surveys, whereas mean group size
and maximum group size for telemetry and circuit surveys did not differ
signiﬁcantly (Table 2.3).

Group composition—The majority of forest buffalo sightings were of >2
individuals (Figure 2.4). Observations of > 2 buffalo usually consisted of several
adult females with their young and one or two adult males that regularly
associated with their respective herds. In 96 circuit survey observations of >2
buffalo, all individuals were identiﬁed to sex and age class. For the 85 sightings
with juveniles and calves, the overall ratio of adults and subadults to juveniles
and calves was 1:2, suggesting that offspring of more than one generation
remain with the females. The ratio of adult males to adult females was 1:6.

Solitary buffalo were more often males (80%) than females (20%; N=102). Of the

49

nine radiocollared females, ﬁve were never observed alone and the other four
collared animals collectively were viewed alone on only nine occasions (AF4 n=4;

AF5 n=2; AF6 n=2; AF8 n=1).

Discussion
Among forest buffalo observations the variation of group size was large (range 1-
46). However, within herds, group size was stable with little variation across
seasons, between savanna and marsh habitat, and time of day. The number of
observations of buffalo increased when the savannas were ﬂush with new grass,
but herd size remained stable. It was also expected that group size would be
larger during morning feeding activity when buffalo use savanna areas (Chapter
I). However, large group size was observed during the morning for only one of
the eight radiocollared buffalo. Also, group size did not differ between marsh and
savanna habitat. Feeding aggregations occurred within herds, but not among
herds.

Herds varied in number of buffalo, with small herds consistently small in
size (i.e. fewer than 10 buffalo) and other herds consistently large with more than
20 buffalo. Forest buffalo group size could be predicted based on previous
observations of the herd, unlike several ungulate species where groupings are
often unstable (Barrette 1991; Clutton-Brock et al. 1982; Estes 1991; Hillman
1987; Leuthold 1970). Herd mates from a single forest buffalo herd were
occasionally separated in different subgroups, but individuals returned to the
herd. Melletti et al. (2007b) observed a similar pattern at Dzanga-Ndkoi NP

where a herd of 24 forest buffalo was often split into two subgroups.

50

Telemetry observations had less bias than circuit survey sightings
because circuit surveys relied on visual observations of buffalo and tended to
miss the smallest groups. Although method of data collection inﬂuences group
size in forest buffalo, radio collars allowed groups of <3 individuals to be
detected, while circuit surveys were useful for monitoring groups that did not
contain a radiocollared buffalo.

Herd size and food resources—Analysis of home range areas indicated a
positive relationship between both maximum group size and mean group size of
forest buffalo with home range area and the amount of open habitat in the home
range area. Further study to determine group size and area of forest clearings
used by buffalo is needed to test this relationship at other forested sites. At other
forested sites in the region, food conditions are similar and buffalo rely on forest
clearings (Blake 2002; Melletti et al. 2007a). Based on results from radiocollared
buffalo at Lopé NP, large group size would be expected for herds that used the
largest clearings.

Forest buffalo social organization—Forest buffalo herds were smaller
than Cape buffalo herds, but were similar in composition. Forest buffalo groups
observed at Lope contained fewer than 50 buffalo, whereas Cape buffalo herds
can exceed 1000 individuals (Sinclair 1977). The core members were adult
females and their young. Unlike Cape buffalo, adult male forest buffalo were
consistently observed with the same herds throughout the year. Taolo (2003)
observed adult male Cape buffalo joining breeding herds in the late dry season at

Chobe National Park, Botswana, and male buffalo alternated between mixed

51

female-young herds and small, male herds at Hluhluwe-iMfolozi Park, South
Africa, during the six-month mating season (Turner et al. 2005). At Lopé, adult
males were not observed leaving herds to form bachelor groups. However, male
forest buffalo were more often observed alone or in pairs than females, and not
all males were associated with a herd.

Solitary aging adult female buffalo were observed at Lopé in addition to
the more common solitary males. For example, three old females (without radio
collars) were often solitary and eventually disappeared. Although no carcasses
were found, these females most likely died before the end of the study. Similarly,
the adult female radiocollared buffalo in the Lopé pilot study (Abernethy 1998,
1999, 2000) and AF4 in this study spent increasingly more time away from their
groups until they died. Instead of becoming solitary, AF6 spent the last four
months of the study associated with a smaller herd. Notably, she was the only
radiocollared buffalo to switch herds. The unusually high number of observations
of aging buffalo was probably due to low predation on adult buffalo and hunting
restrictions. Forest buffalo were one of the main prey items found in leopard scat
at Lope; however, based on analyses of bones in the leopard scat, leopards were
found to prey only on juveniles (Henschel et al. 2005). Due to the low predation
pressure on adult buffalo and enforced hunting restrictions in the study area, food
resources probably have more inﬂuence than predation on group size in forest

buffalo at Lopé NP.

52

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53

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Mean group size

       

ans ‘2’, .-
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AF60 AF8 AF4 AF7 AF5 AF6p AF1 AF3 AF2
(AF9)

Animal lD

 

Figure 2.2. Mean group size of forest buffalo sightings that contained a
radiocollared adult female (AF) at Lopé National Park, Gabon, December 2002-
December 2004. N=the number of observations, representing all observations of
radiocollared buffalo including sightings on circuit surveys and telemetry and
opportunistic observations. Observations include all age classes. Group size
range for each buffalo: AF6c (1-8); AF8 (2-12); AF4 (1-18); AF5 (1-25); AF6p (1-
30); AF1 (2-29); AF3 (5-24); and AF2 (6-46).

 

a)

 

 

b) 50 ’1 AF2
g 40 ‘ r”
0 Ar-jxx
a 30 - AF7 , 0’

c,- AF; AF6p
E 20 ,,/’;”
0 AF3/x ’ AF4
g 10 I ;
AF6c r2 = 0.8706
0 l ‘ i *7 *7 i V 'i 7 i 7 7 'l 7 i
1 2 3 4
Area (kmz) open habitat

 

 

 

 

 

 

Figure 2.3. a) Home range of each radiocollared buffalo in the study area at
Lopé National Park, Gabon (December 2002 - December 2004); b) Maximum
size of group based on observations that included a radiocollared adult female
(AF) forest buffalo in relation to the area of open habitat in the home range of
those females. Each data point represents maximum group size (N=8) deﬁned by
the presence of a radiocollared buffalo. Positive correlation between maximum

group size and areas of open habitat is signiﬁcant (t=6.35, P<0.05).

55

 

 

 

Mean n umber of sightings/circuit

Sep-N 0v

wet

 

Figure 2.4. Seasonal distribution of forest buffalo group size at Lopé National
Park, Gabon, based on circuit survey data. The y-axis represents the number of
observations in each group size class divided by the total number of circuits per
season. Mean group size for all sightings of buffalo, including solitary buffalo,

 

 

 

 

 

Group size class
D1 l2 I>2

_l:l_..

Dec-Feb Mar-May

 

dry
Season

wet

 

  

Jun-Aug

(W

was 1110.34 based on 775 observations during 249 circuits.

56

 

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57

Table 2.2. Correlation coefﬁcients (r) between mean and maximum size of group
and home range characteristics based on adult female buffalo with a radio collar
(N=8).

 

 

 

 

Variable Mean Maximum
group size grog) size

Home range area (kmz) 069* 0.84*

Area of open habitat in home range (kmz) 083* 0.93*

Percent of total home range area that was 0.25 0.09

open habitat

*P<0.05

58

Table 2.3. Comparison of mean group size estimates for telemetry data versus
data collected on circuit surveys for nine radiocollared adult female buffalo at
Lopé National Park Gabon, December 2002-December 2004.

 

 

Group size Mean :1: SE (rye) Mann-Whitney-U test
telemetry circuit surveys

Minimum 2 :l: 0.29 (1-3) 4 :l: 1 (2-8) U=69.5, P=0.0134

Mean 13 :t 2 (4-27) 15 j: 2 (4-25) U=50, P=0.4015

Maximum 24 :l: 4 (8-46) 22 :l: 4 (5-43) U=47, P=0.5660

 

 

59

Chapter III

Korte, L. 2008. Calving and inter-birth intervals of forest buffalo at Lope National
Park, Gabon. African Journal of Ecology.

60

Chapter III

CALVING AND INTER-BIRTH INTERVALS OF FOREST BUFFALO AT LOPE
NATIONAL PARK, GABON

Introduction
The ﬁve subspecies of African buffalo Syncerus caﬁ'er (Sparrman 1779) can be
assigned to the nominate caffer division and the nanus division (Wilson & Reeder
2005). The larger Cape buffalo, Syncerus caffer caffer, (400 to 800 kg) occurs
throughout the savanna areas of eastern and southern Africa, and has a mean
group size of 350 (Haltenorth & Diller 1980; Prins 1996; Sinclair 1977). The
smaller forest-dwelling Syncerus caffer nanus (250-320 kg) is found in the forests
of western and central Africa. Cape buffalo have been the focus of several
comprehensive studies (Mloszewski 1983; Prins 1996; Sinclair 1977); however,
few data exist for forest buffalo (Blake 2002; Melletti et al. 2007a). Based on a
two-year telemetry study, I present calving and inter—birth interval data for forest

buffalo in Gabon and compare results to calving intervals of Cape buffalo.

Materials and Methods
Eight adult female forest buffalo were ﬁtted with radiocollars (Telonics, lnc.,
Mesa, AZ and HABIT, lnc., Victoria, British Columbia) at Lopé National Park, in
December 2002 (AF1-AF8). Due to collar failure, a ninth collar was placed in
December 2003 (AF9). Animals were tracked on a weekly basis (mean number
of locations = 205/buffalo) from December 2002 through December 2004. When
radiocollared animals were observed, notes on young in close proximity to adult

females were recorded (i.e. age, sex, distinct characteristics). A Sony digital

61

video camera (model DCR-TRV140) was used to record mother-young pairs
when the view was clear. Collared females were from eight different herds having
deﬁned home ranges with small areas of overlap (Chapter I).

Buffalo home ranges were predominately savanna habitat despite greater
availability of forest habitat at the study site (Chapter I). A controlled burning
program maintains these savannas, creating open areas for tourists to observe
large mammals (White & Abernethy 1997). Savannas, burned during the June
through August dry season, provide a ﬂush of new grasses grazed by buffalo
during the September through November wet season. Green grass, available
during the December through February dry season, becomes tall, dry, and less

palatable for buffalo by the March through April wet season.

Results
Four collared females (AF1, AF2, AF3, AF8) had a single young with them when
collared, and their young remained in close proximity to them throughout the two
years of study (Table 3.1). Of those, three did not calve after collaring. The fourth
(AF8) had a second calf when her ﬁrst calf was two years old. The ﬁrst calf
remained with her after the birth of the second calf.

Four collared females (AF4, AF6, AF7, AF9) provided little information on
calving intervals. AF4 was never observed with young. Each of the remaining
three had no young at the time of collaring, but had a single calf during the study.
AF6 had a calf four months after collaring in April 2003, but lost the calf by July
2003. She was not observed with another calf through the end of the study. AF 7

had a calf 19 months after collaring in July 2004, but lost the calf within four

62

months. AF9 was collared in December 2003, had a calf in August 2004, and lost
the calf within a month. AF7 and AF9 lost their calves during the last four months
of study, leaving insufﬁcient time for gestation of another calf.

The remaining radiocollared animal (AF5) gave birth twice during the two-
year study. She calved in March 2003, but lost her calf within four months. She
was observed mating with an adult male on 19 August 2003 and her second calf
was born in June 2004, suggesting a calving interval of 16 months. The
estimated 10 month gestation period is similar to the 11.5 gestation period
reported for Cape buffalo (Sinclair 1977). Her second calf survived through the
remaining six months of study.

In summary, ten young were observed with the nine radio-collared
females. Four of those were with radiocollared animals at the time of collaring
and remained with the females throughout the two-year study. Six were born
after collaring. Two of those calves survived the remaining six months of study.
Four of the six calves disappeared before reaching four months of age (i.e., 67%

mortality).

Discussion
The calving interval at Lopé is at least two years between surviving young.
Similar intervals are reported for Cape buffalo with a range from 15 to 25 months
at sites in South Africa, Tanzania, and Uganda (Grimsdell 1969; Prins 1996;
Sinclair 1977). Because food quality improved with grazing pressure, early work
suggested a positive relationship between population density and reproductive

rate in African buffalo (Grimsdell 1969); however, a long calving interval (two

63

years) has been recorded for Cape buffalo in Tanzania (Prins 1996) despite a
high population density (18-20 buffalo/kmz). In this case, social factors and herd
structure also inﬂuenced reproductive rate, indicating that population density
alone does not explain calving intervals of Cape buffalo (Prins 1996). At Lopé,
forest buffalo exhibit a relatively long calving interval with a low population
density (~5 buffalo/kmz; Chapter II). Clearly, there is a need to further investigate
factors that inﬂuence calving intervals.

This study found a high mortality rate for calves (67%). At Lope, cause of
death for the calves was not determined because carcasses were not found.
However, buffalo calves are a primary prey of leopard Panthera pardus in the
study area (Henschel et al. 2005). Another important factor may be food
availability. The two surviving calves born in June and July, and their lactating
mothers soon had access to the seasonal ﬂush of new grass during the
September through November wet season. The two calves born in March and
April when food resources were low did not survive past four months. In Cape
buffalo, breeding ecology is closely related to seasonal availability of resources
(Pienaar 1969; Prins 1996; Ryan et al. 2007). The long calving interval and high
calf mortality at Lopé may contribute to a low reproductive rate. Further research
of the reproductive biology of forest buffalo, especially in regard to the

seasonality and availability of resources, is recommended.

64

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65

Chapter IV

HERD-SWITCHING IN ADULT FEMALE AFRICAN FOREST BUFFALO
SYNCERUS CAFFER NANUS

Introduction
Recent evidence suggests that herd-switching in African buffalo Syncerus caffer
(Sparrman 1779) is more common than previously reported (Cross et al. 2005;
Halley et al. 2002). Traditionally, buffalo herds were considered stable with little
movement of buffalo among herds (Estes 1991; Kingdon 1982; Prins 1996;
Sinclair 1977). Although male buffalo were known to form small bachelor groups
when not with the large herds, female buffalo and their young were viewed as
core herd members. Recent studies in Botswana and South Africa have
documented adult females switching herds and dispersing distances of > 100 km
(Cross et al. 2005; Halley et al. 2002). These studies focus on Cape buffalo,
Syncerus caffer caffer, a savanna-dwelling subspecies; however, less is known
about the smaller, forest-dwelling subspecies, Syncerus caffer nanus. | report a
single case of herd-switching over a distance of < 1 km from a telemetry study of

nine adult female forest buffalo at Lopé National Park, Gabon.

Material and Methods
In December 2002, eight adult female forest buffalo, representing eight different
groups, were ﬁtted with radio collars (Telonics, Inc., Mesa, AZ and HABIT, Inc.,
Victoria, British Columbia) at Lope National Park. Due to collar failure, a ninth
collar was placed in December 2003. This ninth female (AF9) was in the same

group as AF5. Buffalo were tracked on a weekly basis from December 2002

66

through December 2004 with a mean total number of locations of 205 per buffalo.
When radiocollared buffalo were observed, notes on all buffalo in the group were
recorded in order to distinguish individual buffalo as well as herds. In addition, a
Sony digital video camera (model DCR-TRV140) was used to record buffalo

when the view was unobstructed.

Results
One radiocollared buffalo switched herds. Between December 2002 and July
2004, she was observed in a herd with a mean group size of 17 (range 4-30). In
August 2004, she joined a smaller herd of four buffalo where she remained the
last ﬁve months of the study. The switch from a larger to a smaller herd, resulted
in a reduction of home range from 4.17 km2 to less than half that (i.e., 1.54 km").

The other eight radiocollared buffalo remained with their respective herds.

Discussion
The single case of herd-switching among nine radiocollared buffalo at Lopé NP
suggests that forest buffalo herds are more stable than Cape buffalo herds.
Among Cape buffalo, herd-switching can be frequent among adult females
(Cross et al., 2005; Halley et al., 2002), but only one radiocollared adult female
switched herds in this study of forest buffalo. To better understand social
networks, Cross et al. (2005) proposed a new association index based on data
from Cape buffalo at Kruger National Park, South Africa. In the Kruger study,
ﬁssion events were deﬁned as two radiocollared animals observed in the same

group and subsequently observed in different groups. For the Lope buffalo, there

67

were no cases of two radiocollared buffalo observed in the same group and
subsequently observed in different groups.

If Lope buffalo had a similar ﬁssion pattern to that of the Kruger buffalo,
then about 19 ﬁssion events would be expected. At Kruger, where approximately
90 Cape buffalo were radio-collared among 4-12 herds with a combined
population of 3,000 buffalo, 185 ﬁssion events were recorded among adult
females and juveniles over a two-year period. Nine collared forest buffalo were
tracked among 18 herds in the Lope population of 342 buffalo. However, no
ﬁssion events occurred among radiocollared buffalo at Lopé and only one
radiocollared buffalo switched herds during the two-year study. In addition to
radiocollared buffalo, the same recognizable individuals (i.e., about 6—10 adult
females and 1-2 adult males per herd) were consistently observed with herds.
Thus, my data suggest that ﬁssion events are less common in forest buffalo than
in Cape buffalo.

Distribution of resources may inﬂuence ﬁssion events in African buffalo. In
Botswana, herd-switching occurred during wet seasons when Cape buffalo herds
migrate and fragment into smaller subgroups (Halley et al. 2002). Water
resources are not limited during the dry season and forest buffalo do not migrate
at Lope NP (Chapter I). Small streams ﬂow throughout the forest habitat and
each herd has several water resources within their home range areas. Forest
buffalo home range areas are well deﬁned with small areas of overlap (Chapter
I). Despite having adjacent home range areas, radiocollared buffalo from different

herds were never observed together.

68

Small herd size may explain the rarity of ﬁssion events observed among
forest buffalo. Cape buffalo group size is considerably larger than that of forest
buffalo at Lopé (i.e., 12 :I: 2 with a range of 3-24; Chapter II). Although groups as
small as 12 have been observed in Cape buffalo, mean size is 350 with the
largest herds reaching into the thousands (Prins 1996; Sinclair 1977). Halley et
al. (2002) suggest that subgroups rather than individuals are switching among
Cape buffalo herds. Cape buffalo herds appear to be aggregations of subgroups,
whereas forest buffalo herds are small and stable with relatively little switching
between herds. Notably, the only buffalo to switch herds was an older buffalo and
she was the single member of her original herd to switch. Her switch may have
been a reﬂection of her condition and inability to move with her original larger
herd. Further investigation of association patterns in forest buffalo with an
increased sample size of collared buffalo would be useful to understand why

buffalo switch herds.

69

Chapter V

PERSPECTIVES ON THE ROLE OF BUFFALO IN CONSERVATION AT LOPE
NATIONAL PARK, GABON

Introduction
Conservation planning for wildlife depends on knowledge of the animal's biology
as well as the management objectives within protected areas. The newly created
parks in the Congo Basin Forest region of Central Africa present an opportunity
to consider fresh approaches to park management. The Congo Basin is a priority
for global conservation and is one of only three regions of the world (along with
the Amazon Basin and the island of New Guinea) designated as a tropical
wilderness area (Mittermeier et al. 1998). These regions are recognized for their
high biodiversity combined with large expanses of intact tropical forest and low
human populations. Though the tropical forest of the Amazon Basin covers a
larger area than the Congo Basin, many locations in the Amazon are devoid of
large animals (Redford 1992). Only the Congo Basin forests retain relatively
abundant populations of large mammals.

The Amazon rainforest and the savannas of Africa have been the focus of
conservation activities for many years, but Africa’s Congo Basin only recently
received international recognition as a conservation priority (Kamdem-Toham et
al. 2003). In the 1999 Yaoundé Declaration, six heads of state from Central
African nations committed 10% of forest areas in their countries to become
protected areas to safeguard their tropical timber and wildlife. The region’s

wildlife includes many charismatic, large mammals such as gorillas (Gorilla g.

70

gorilla), chimpanzees (Pan t. troglodytes), mandrills (Mandn'llus sphinx),
elephants (Loxodonta cyclotis), and buffalo (Syncerus caffer nanus). In 2002,
President Omar Bongo of Gabon fulﬁlled his promise by creating thirteen new
national parks covering 30,000 km2 or 10% of Gabon’s land mass. At 5,000 kmz,
Lope National Park is one of the largest of the parks and the conservation
community views it as the emerging Yellowstone of Gabon.

Though the creation of national parks provides the legal documentation to
protect habitats, laws alone will not lead to conservation of natural resources
(Brashares et al. 2001; Newmark & Hough 2000). Efﬁcient and well-planned
activities are essential for conservation, and fundamental ecological data are
crucial for planning (Milner-Gulland & Bennett 2003). However, planning in the
Congo Basin is difﬁcult because data on animals, plants, and social conditions
are often lacking, and protected areas are often ineffectively managed (Blom
2004; Kamdem-Toham et al. 2003). Further research can provide information for
planning and zoologists must be proactive in making recommendations that will
contribute to the success of park management plans (Pullin & Knight 2001).
However, to build the capacity of African wildlife biologists, greater efforts are
needed to integrate research, training, and park management (Durant et al.
2007; McNeilage & Robbins 2007). The case study of the Serengeti Cheetah
Project by Durant et al. demonstrates successful integration of long-term
research and management in conservation. In this ﬁnal chapter, I consider how
long-term research of forest buffalo could be integrated with park management

and training at Lope NP.

71

Throughout my two-year study on the spatial and social organization of
forest buffalo at Lopé National Park (Chapters l-lV), I had the opportunity to
conduct research, collaborate with park managers, and mentor national and
international students. Park management activities (i.e., savanna burning)
inﬂuence buffalo behavior (Molloy 1997), buffalo provide relatively easy viewing
for park visitors, and buffalo have been a focal species for training of national
wildlife professionals. Given the important role buffalo play in this region, an
increased effort should be made to expand conservation activities at Lope NP to
include this species. Such a program must ﬁt with other conservation priorities
given limited personnel, funding, and expertise. However, a focus on forest
buffalo that integrates research, park management, and training could make a
positive contribution to conservation planning. After an introduction to African
buffalo and the institutions at Lopé NP, I consider each of the objectives of the
Lope National Park Management Plan for 2006-2011 and discuss how further
research of forest buffalo could contribute to achieving each objective.
Regionally, establishment of a long-term conservation study of buffalo could play

a leading role in making Lopé NP a model for conservation in the Congo Basin.

Forest buffalo
Forest buffalo are one of ﬁve recognized subspecies of African buffalo (Wilson &
Reeder 2005). These subspecies differ in geographic range, morphology, and
behavior. The largest of the subspecies is Syncerus caffer caffer, the 400-800 kg
Cape buffalo of the savannas of east and southern Africa (Grubb 1972; Kingdon

1997; Sinclair 1977). At 250-320 kg, forest buffalo of the Congo Basin are the

72

smallest of the subspecies (Haltenorth & Diller 1980; Kingdon 1997). Cape
buffalo also have largest groups, with a mean group size of 350 and large herds
that reach into the thousands (Prins 1996; Sinclair 1977). Among forest buffalo,
group size is small with a mean group size of less than 20 individuals (Chapter 2;
Jarrnan 1974; Melletti et al. 2007b; Mloszewski 1983; Sidney 1965). Cape buffalo
are known for the distinct lateral curve of their horns, whereas forest buffalo lack
the lateral extension and have small, swept back horns. Kingdon (1982) suggests
that this suite of differences observed among the African buffalo subspecies
reﬂect adaptations for living in savanna versus forest habitat.

Forest buffalo are recognized as an important species for conservation
planning based on their spatial requirements, ecological function, vulnerability,
and socioeconomic signiﬁcance (Blake 2002; Coppolillo et al. 2004; Melletti et al.
2007a). At Lope NP, buffalo home range areas are predominately savanna
habitat, but buffalo also use forest and marsh habitat within these ranges
(Chapter I). Thus, their home ranges and use of these areas include all habitat
types at Lope NP. Buffalo are the only large grazing ungulate and juvenile buffalo
are an important prey species for leopards (Panthera pardus, Henschel et al.
2005). Though protected near park headquarters, buffalo are vulnerable to
hunting in Gabon (Laurance et al. 2006b). People value buffalo for their meat as
well as their potential as one species in a suite of African forest mammals that
will draw international tourists. Working with local communities and developing

tourism are both objectives of the Lopé NP Management Plan.

73

Lopé National Park Management Plan
With the enthusiasm generated by the creation of the new national parks in
Gabon, researchers and park staff created the Lopé National Park Management
Plan for 2006-2011. The long-term vision of the plan is to have Lopé NP become
a model and leader for conservation efforts in the Congo Basin. The
management plan includes a desire to incorporate local, national, and
international interests and values in its management objectives. Clearly, this will
be a major challenge since expectations for how the park resources are used
differ among these stakeholders. For example, when international tourists visit
Lopé NP, they hope to see as many large mammals as possible. In contrast,
local communities prefer low densities of wildlife to reduce the risk of large
mammal confrontations, including crop-raiding and threats to people. A focus on
buffalo has the potential to manage the diverse expectations for Lopé NP and

surrounding communities.

Park management, research, and training institutions at Lopé National Park
The park headquarters, the Lope research center, and the Lopé Training Center,
are based within 11 km of each other. The park headquarters is supported by the
Gabonese government and there is a history of signiﬁcant funding from the
European Union. The history of legal protection of the region began in 1946
when the area was designated the Lopé—Okanda Wildlife Reserve. In 2002, the
Lopé—Okanda Wildlife Reserve became Lope National Park with the creation of

the 13 national parks in Gabon.

74

Researchers based at the Lope research station are advisors to park
management and work closely with park staff to plan activities, such as savanna
burning and monitoring of wildlife populations within the park. The Gabonese
lntemational Center for Medical Research (CIRMF) founded the Lopé research
station (Station d’Etudes des Gorilles et Chimpanzés, SEGC) for the study of
gorillas and chimpanzees in 1983. Since the 1990s, the research station has
expanded its program to include the general ecology of the park. The Lope
research station has the longest running continuous ecological monitoring
research program in the region, representing a valuable source of data for
comparative and longitudinal studies. In addition, the Wildlife Conservation
Society (VVCS) provides the Lopé research station with funding for conservation
activities. In 2003, the director of the Lopé research station established the Lope
Training Center to improve scientiﬁc capacity of Gabonese conservation

professionals.

History of buffalo research at Lopé
Research on Lope forest buffalo began in the late 1980s and continues today.
The ﬁrst buffalo study focused on the vegetation of the savanna areas (Alers &
Blom 1988). Field surveys of mammals conducted in the 1990s documented the
abundance of forest buffalo in the savanna areas (T utin et al. 1997), but found
few signs of buffalo in forest habitat (White 1994). Following those surveys,
Molloy (1997) examined the inﬂuence of savanna burning on forest buffalo. In
1998, radio-collars were placed on one adult female and one adult male to test

the feasibility of a telemetry study of buffalo at Lope NP (Abernethy 1998, 1999,

75

2000). Subsequently, a two-year telemetry study during 2002-2004 based on
nine radio-collared buffalo representing eight different herds examined the spatial
and social organization of Lopé buffalo (Chapters l-IV). Van Hooft et al. (2000)
used tissue samples from Lope buffalo for a study of African buffalo population
genetics. Most recently (2006), Van Hooft collected additional genetic samples,
and during the summer of 2007, one of his graduate students collected forest
buffalo dung samples for a preliminary study of buffalo diet and parasites.
Although many species at Lope NP might serve as a focus for
conservation efforts, buffalo have characteristics that make them especially
suitable. First, conditions for buffalo research at Lope NP are favorable with low
predation, little hunting pressure, and seasonally abundant food resources
(Chapters l & ll). Second, forest buffalo research is straightfonrvard compared
with other sites in the Congo Basin. At other sites, buffalo tend to use forest
clearings that are scattered throughout difﬁcult terrain with typically only one herd
using each clearing or set of clearings (Blake 2002; Melletti et al. 2007b). In
contrast, several herds of buffalo can be studied simultaneously in the savanna
areas at Lope NP. These savanna areas also facilitate telemetry studies because
the buffalo can be collared at minimal risk to the buffalo and buffalo can be
tracked easily. In addition, buffalo are clearly visible in the forest-savanna habitat

mosaic after savanna burning (Molloy 1997).

Lopé NP Management Plan Objectives
Protect and conserve species.—The ﬁrst objective of the Lope NP Management

Plan is to protect species. All populations of African buffalo (Syncerus caffer) are

76

categorized as lower risk/conservation dependent on the lUCN Red List of
Threatened Species (lUCN 2007). Being a conservation dependent species
means that if existing conservation activities for buffalo were to end, the African
buffalo would be listed in a threatened category, a more severe listing. However,
this assessment is out-of date and the population trend was downward at the last
assessment. Given that African forests are threatened by logging, hunting of
wildlife, and insufﬁcient funds to manage protected areas (Kamdem-Toham et al.
2003), there is a need to protect and conserve forest buffalo. A long-term
conservation study which collects biological, behavioral, ecological, and health
data of importance to the conservation of African buffalo in the Congo Basin
would provide managers of protected areas with data to inform their decision
making.

Conserve habitat and ecological processes—The Lopé NP Management
Plan calls for the conservation of unique habitats and ecological processes within
the park. The majority of Lopé NP is forest and most importantly the park
protects tropical forest habitat. However, the northeast corner of the park is a
mosaic of forest-savanna habitat, which has also been noted as an important
habitat to protect and sustain. This mosaic supports abundant wildlife populations
and a unique array of plant species (T utin et al. 1997; Ukizintambara et al. 2007).
The savanna areas were traditionally maintained by early inhabitants of Lope and
today park managers preserve open habitat through savanna burning to create

scenic vistas for tourists (Tutin et al. 1997; White & Abernethy 1997). Without the

77

annual savanna burning, the savannas areas would be colonized by forest
species and return to forest habitat (Oslisly 2001).

Forest buffalo include savanna area within their home ranges and use the
forest-savanna mosaic with no migration into the continuous forest (Chapter I).
After the savanna burning during the June through August dry season, buffalo
feed on the ﬂush of new grass from September through February and use the
forest habitat adjacent to the savannas during March through August when the
savanna grasses are dry, tall, and less palatable. Home ranges of forest buffalo
are stable in size and location over at least a few years and might remain stable
over several years (Chapter I). Given the close link between savanna condition
and forest buffalo use of savanna habitat, buffalo are a key species for guiding
savanna management.

Traditionally, savannas are burned by igniting savanna edges and
allowing large swaths of savanna to burn. These areas are burned over a few
days in late August and early September, creating one or two large ﬂushes of
new grass. To create additional ﬂushes of grass over longer time periods, I
suggest that small burns are staggered in time and space within buffalo home
range areas (Figure 1.2). Within home ranges, ﬁrebreaks could be used to limit
ﬁres to discrete patches. Burning savannas over a longer time period could
provide buffalo herds with new grass over several weeks and increase the period
of buffalo visibility, a beneﬁt for tourism as well as research and training. The

greater number of ﬂushes may also lead to an increase in the number of buffalo

78

given the additional food resources. These beneﬁts will be discussed further in
the following sections, which address tourism, research, and training.

A potential problem with multiple burning is the amount of time and labor it
takes to implement this plan. Based on my ﬁeld assistant’s preliminary trials of
this method, burning in small patches over several months is a more laborious
and time-consuming process than the currently used method of burning large
savanna areas over a short time period without ﬁrebreaks. Therefore,
professional training in prescribed burns is recommended for park managers as
well as logistical support and incentives to implement plans. After the savanna
burning, the time period buffalo use savannas and number of buffalo using the
savannas should be monitored to determine how the burning patterns inﬂuence
buffalo. Further study should not be limited to buffalo. The impact of savanna
burning on other species including plants and animals has yet to be assessed.
Thus, further study of the impact of savanna burning will be needed to evaluate
how well habitats are conserved and ecological processes are maintained at
Lope NP.

Protecting archeological sites.—To my knowledge, forest buffalo and
archeologists have yet to clash at the archeological sites scattered throughout
the Lopé savannas. However, buffalo often rest on hilltops that include
archeological sites. From the hilltops, there are clear views across the savanna
to the forest edge and often a breeze to ward off heat and insects. And, the Lope
savannas support high densities of elephants and buffalo (Tutin et al. 1997).

Lopé NP contains archeological sites dating from the Upper Pleistocene to the

79

Post-colonial period and the excavated sites at Lopé cover the longest and most
complete archeological sequence in Central Africa (Ndong 2002). The 40,000
years of human presence in the region contributed to maintaining the forest-
savanna mosaic at Lope (Ndong 2002; Oslisly 2001). Thus, the Lope sites
represent a long history and need adequate protection. The Lopé NP
Management Plan recommends that more delicate and older archeological sites
may need to be sheltered from high concentrations of wildlife. Buffalo use
archeological sites, but their impact on these sites has yet to be investigated.
Therefore, buffalo use of archeological sites should be quantiﬁed to reduce the
risk of any potential damage to important sites by buffalo.

Developing proﬁtable tourism activities.—A long-term study of forest
buffalo could beneﬁt tourism activities. In particular, a well-implemented savanna
burning plan could result in several buffalo herds clearly visible in the savanna
areas over extended periods of time. Seeing buffalo is exciting for tourists,
especially when other wildlife in the area less visible. Buffalo on the savannas
also assure visitors that the wildlife in the park is well-protected. When tourists
drive through empty savanna areas without buffalo, it can be difﬁcult for them to
appreciate why Lopé is a national park. The creation of the national parks was
expected to generate alternate sources of revenue, and Gabon needs to diversify
its economy (Laurance et al. 2006a). Though gorillas are what attract tourists to
Gabon, savannas must be populated with wildlife for tourism to succeed. Tourists
visiting Africa expect to see abundant wildlife, especially if they have previously

visited sites in eastern and southern Africa. Therefore, Gabon will need to

80

maximize the potential of each wildlife encounter. Tourists will gain an
appreciation for the abundance of wildlife at Lope NP, when they take a sunset
drive on a savanna among several herds of buffalo.

If park revenue can be generated from forest buffalo, it would reduce
dependency on international funding and base economic gain in local resources
and institutions. When appropriately managed, safari hunting generates revenue
and contributes to community development (Lewis 1995; Loveridge et al. 2007;
Mayaka et al. 2005). Although hunting is illegal at Lopé NP, it may be worth
considering the participation of international sport hunters in the darting of buffalo
for radio collar placement. Another consideration would be to involve sport
hunters in the handling of problem buffalo. On occasion, park management will
kill a buffalo if a particular buffalo has injured a person. However, a clear and
consistent protocol for addressing problem buffalo does not appear to exist.
Having sport hunters involved could potentially increase revenue for the park via
trophy fees as well as reduce the negative impact of wildlife on the local
community.

Promoting local participation in all conservation activities—Positive
interactions between members of the Lopé community and wildlife could increase
the community’s interest in protecting species and supporting the park. The
increased visibility of buffalo on the savanna can also be used to provide the
local communities with opportunities to view wildlife. Forest buffalo are an ideal
species to engage the local community because buffalo are relatively easy to

observe compared with other African forest mammals. Tracking of radiocollared

81

buffalo provides a thrilling wildlife demonstration and often leads to visual
observations of the collared buffalo. Positive interactions with animals give
people a better understanding and appreciation for wildlife (Moorman 2006).
When I took school children to radio-track buffalo, they could follow a signal from
the radio collar and eventually identify the collared buffalo in a herd. Forest
buffalo were rather undisturbed by this interaction and several children could
participate. Buffalo will continue to feed, rest in a wallow, or meander across the
savanna despite children’s observations. Positive interactions between members
of the local community and wildlife will increase the community’s interest in
protecting species and supporting the park (Steinmetz et al. 2006).

The success of any park will depend on the park having a positive impact
on the local community (Wilkie et al. 2006). Beneﬁts to the local community must
come through employment, development, or activities managed by park staff
because people are prohibited from harvesting natural resources in the park.
Though some employment opportunities exist for tour guides and work with the
Lopé research station and training center, these positions beneﬁt only a few
individuals. Funding from the European Union has helped to support
infrastructure for the park, but is neither consistent nor unlimited. The park has
not generated sufﬁcient revenue to provide funding for development projects
such as building schools or clinics. It seems the local community would have
greater incentive to reduce poaching if they received direct beneﬁts from the

protected wildlife.

82

Managing forest buffalo for a sustainable harvest to beneﬁt the local
community could be considered since buffalo populations appear to be
increasing. The number of herds and individual buffalo at Lopé increased
between 1996 and 2004. l estimated that 212 buffalo in 13 herds used the study
area between July and December 1996 (Molloy 1997). During the 2002-2004
study, the estimated population was 342 buffalo in 18 herds (Chapter II). This
increase is probably due to low predation on adult buffalo and the enforcement of
hunting laws. Although Henschel at al. (2005) found evidence of leopard
predation on juvenile buffalo in the study area, they found no evidence of
predation on adult buffalo. The frequent observation of older solitary females at
Lope NP suggests that predation pressure on adult buffalo is low (Chapter II). In
addition, the Lope buffalo population is protected from hunting due to the
proximity to park headquarters. In southwest Gabon, areas where hunting
pressure is high have fewer forest buffalo than sites with no hunting (Laurance et
al. 2006b).

The consideration of sustainable harvest of buffalo should not be done at
the expense of the good viewing conditions for buffalo at Lopé NP. To maintain
high numbers of buffalo and calm herds, a transparent system for determining
the number of buffalo in the population is required. And, the appropriate number
as well as age and sex class to harvest over what time period would need to be
determined.

Develop training and multidisciplinary research—The ﬁnal objective of the

Lopé NP Management Plan is to have the park become a resource for training

83

and research in the Congo Basin. An institutional collaboration between the
research station and the training center to establish a long-term conservation
study of forest buffalo would advance scientiﬁc research and contribute to park
management and capacity building of national scientists. Few studies have
focused on African forest mammals and there is still much to be learned about
the behavior and ecology of forest buffalo. As previously discussed, conditions
for studying buffalo at Lopé NP are favorable with the ability to access and study
several different herds. Collaboration among park management, the research
station, and the training center can be facilitated by their geographic proximity
and current cooperative relationships. Since the establishment of the training
center, trainees have participated in buffalo research and learned telemetry
techniques by tracking forest buffalo. Buffalo proved to be an ideal species for
learning about animal behavior and survey techniques because buffalo can be
readily observed from a distance and radio-tracked with little disturbance of the
herds. From a multidisciplinary perceptive, study of the savanna burning at Lope
NP is an excellent opportunity to link studies of vegetation with buffalo biology

and park management activities.

Summary of recommendations
This paper took a ﬁrst look at how each management objective of the Lope
National Park Management Plan relates to forest buffalo. Ultimately, managers
must determine whether they would like wildlife populations to increase,
decrease, or remain stable, and to implement appropriate management plans.

Table 5.1 is a summary of the objectives of the Lopé NP Management Plan with

84

the management recommendations and potential beneﬁts that could result with
implementation of these suggestions. Clearly, further work will be needed to
determine how any one of these recommendations could be implemented. And,
forest buffalo would need to be monitored to provide the data for appropriate
planning. For example, home range area estimates should be updated on an
annual basis for planning savanna burns. Though harvest of buffalo could be
considered to beneﬁt the local communities, it would require detailed population
data to determine appropriate harvest levels as well as the time periods between
harvests. Given buffalo data required to make informed management decisions,
a long-term conservation study on forest buffalo could make a signiﬁcant
contribution to park management. In addition, a long-term conservation study will
provide opportunities for national scientists to develop their ﬁeld research skills.
Buffalo have the potential to be a model species for integration of research,

management, and capacity building at Lopé National Park.

85

 

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86

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