Detecting Changes in Elephant Body Condition in Relation To

Detecting changes in elephant body
condition in relation to
resource quality
By
Christelle de Klerk
Submitted in fulfillment
of the requirements
for the degree of
Magister Scientiae
in the Faculty of Science
at the
Nelson Mandela Metropolitan University
Supervisor: Prof G.I.H. Kerley
January 2009
ACKNOWLEDGEMENTS
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This project was made possible through funding by the National Research
Foundation (NRF), a NRF grantholders bursary through Prof. Graham Kerley,
and NMMU postgraduate bursary.
I’d like to thank my supervisor, Prof. Graham Kerley, for the opportunity to work
on this project. Thank you for all your input, guidance and support throughout, as
well as the time spent reading drafts of this document.
I’d also like to thank all the owners, managers and staff of the various game
reserves for always being ready to accommodate and assist me with my project.
Specifically, I’d like to thank John O’Brien and Bruce Main from Shamwari,
Warrick Barnard from Blaauwbosch, Jennifer and Mike Fuller from Kariega, and
Richard and Kitty Viljoen from Asante Sana for always welcoming me and going
out of their way to make sure I had everything I needed. It was a fantastic
experience. I’d also like to thank SANParks for the opportunity to work in the
Addo Elephant National Park.
My sincerest thanks to Marietjie Landman and Craig Tambling for assistance with
my statistics. I would definitely have been very lost indeed without your help!
I’d also like to thank my two field assistants, Alexander Finger and Francesca
Lyndon-Gee, for their company, friendship and help during my field trips. You
guys just made the whole experience that much better.
Lastly, I’d like to thank all my friends and family who embarked on this journey
with me. Thank you for your understanding, support and encouragement
throughout this entire project. I would not have been able to do it without you.
A special thank you to my mom, dad and Philip- your continued encouragement,
love and belief in me is what got me this far.
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CONTENTS
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Acknowledgements i
Contents ii
Abstract v
Chapter 1
General introduction
1.1 Introduction 1
1.2 The elephant problem 1
1.3 Elephant life history and population regulation 4
1.4 Caughley’s framework 6
1.5 Body condition as indicator of response 7
1.6 Elephants as herbivores 7
1.7 Measuring diet quality 8
1.8 The elephant problem in the Eastern Cape 8
1.9 Rationale, objectives and research approach 9
Chapter 2
General description of the study sites and populations
2.1 Background and elephants in the Eastern Cape 11
2.2.1 Historic populations and distribution 11
2.2.2 Decline of the elephant 13
2.2.3 Current populations and distribution 14
2.2 Study sites and populations 15
2.2.1 Addo Elephant National Park- Main Camp 15
2.2.1.1 Site description, topography and geology 15
2.2.1.2 Climate 16
2.2.1.3 Vegetation 17
2.2.1.4 Elephant population history 18
2.2.1.5 Elephant diet 18
2.2.2 Addo Elephant National Park- Nyathi Concession Area 19
2.2.2.2 Climate 19
2.2.3 Asante Sana Private Game Reserve 21
2.2.3.2 Climate 22
2.2.4 Shamwari Private Game Reserve 24
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2.2.4.2 Climate 25
2.2.5 Kariega Private Game Reserve 27
2.2.5.2 Climate 28
2.2.6 Blaauwbosch Private Game Reserve 31
2.2.6.2 Climate 31
Chapter 3
Life history variation in body condition
3.1 Introduction 34
3.1.1 Body condition 34
3.1.2 Body condition over lifetime 34
3.1.3 Condition measures 35
3.2 Hypotheses and aims 36
3.3 Materials and methods 37
3.3.1 Selection of life-history groups 37
3.3.2 Data collection 38
3.3.3 Data analysis 40
3.4 Results 40
3.5 Discussion 45
3.5.1 Condition and life stage 45
3.5.2 Condition and season 48
Chapter 4
Site-specific variation on body condition
4.1 Introduction 50
4.1.1 Spatial variation in forage resources 50
4.1.2 Impacts of herbivores on forage resources 50
4.1.3 Effects on body condition 52
4.1.4 Diet quality and its assessment 52
4.1.5 Elephant nutrient requirements 55
4.2 Hypotheses and aims 56
4.3 Materials and methods 56
4.3.1 Choice of sampling method 56
4.3.2 Measures of dietary quality 57
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4.3.3 Sampling procedure 58
4.3.4 Data analysis 59
4.3.4.1 Body condition 59
4.3.4.2 Dietary quality 60
4.4 Results 61
4.4.1 Body condition 61
4.4.2 Dietary quality 74
4.4.2.1 Precision of sample analysis 74
4.4.2.2 Faecal dietary quality 74
4.4.2.3 Plant dietary quality 80
4.5 Discussion 81
4.5.1 Population specific patterns 81
4.5.2 Dietary quality variation 87
Chapter 5
Concluding discussion
5.1 Body condition scores 91
5.2 Responses in body condition 91
5.3 Future Research 94
References 97
Appendix 1 – Sample sizes obtained for the various sites, seasons and
life stage stages 111
Appendix 2 – Results of the log-linear analysis of body condition and
life stage in the Addo Main Camp 112
Appendix 3 – Results of log-linear analysis of body condition for all
sites and seasons 113
Appendix 4 – Results of the multinomial regression model 114
Appendix 5 – Results of the ANOVAs for faecal dietary quality 115
Appendix 6 – Plant dietary quality (protein, phosphorus and NDF)
values and results of overlapping plant species and
PDI’s 116
Appendix 7 – Rainfall recorded at each site for 2007/2008 119
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ABSTRACT
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Elephants, as megaherbivores, are known to have extensive impacts on
vegetation, especially in enclosed areas. This raises the issue that elephants in
enclosed areas may become limited by resource availability. Resource limitation
is generally expressed via density dependence, but elephants, due to their slow
demography, may not be affected by initial changes in resource availability. This
highlights the need for a more sensitive measure of resource limitation to allow
for the detection of energy stress within a population before changes in vital rates
occur. This study investigated visual changes in elephant body condition in
relation to resource availability in a number of Eastern Cape reserves to assess
whether body condition could be used to detect life stages, as well as seasons
and sites which may be resource limited. Elephant life stages were divided into
energy stressed (newly weaned calves, lactating females, and old females) and
non-energy stressed classes (sub-adults and non-lactating females) to determine
whether energy stressed life stages were more vulnerable to resource limitation.
In the AENP it was found that lactating and old females exhibited significantly
poorer body condition than non-energy stressed individuals, but that weaned
calves had body conditions similar to non-energy stressed individuals.
Comparisons between seasons revealed that all life stages exhibited better
condition in winter than summer or spring, with lactating females showing little
recovery of condition over time. Seasonal body conditions were correlated with
rainfall recorded in the Addo Elephant National Park. Comparisons of elephant
body condition between sites (n = 6) revealed that body condition varied across
sites, with poorer body condition associated with areas of higher elephant density
and low rainfall during the study period. Comparisons with faecal dietary quality
data both between sites and seasons indicated that body condition also
responded to changes in the availability of protein and neutral detergent fibre
(NDF) of plant resources, with higher protein and lower values associated with
better condition. Based on condition estimates of elephants occurring in the Addo
Main Camp, it was established that this population is experiencing nutritional
stress, with energy stressed individuals exhibiting the lowest body conditions.
This was supported by dietary quality measures. Our findings suggest that
elephant body condition is a good measure for detecting resource limitation, both
within populations and between seasons, and that elephant body condition
respond to relatively small changes in resource conditions, thus making it an
effective measure for the detection of nutritional stress. Additionally, our findings
show that energy stressed individuals, particularly lactating and old females are
more vulnerable to resource limitation. This demonstrates the importance of
monitoring these life stages for the detection of density dependence within
populations. Finally, our data suggest that threshold values of faecal dietary
quality may exist at which body condition within a population begins to
deteriorate, making it possible to determine the condition of a population through
values obtained in faecal samples.
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Keywords: Body condition, dietary quality, protein, phosphorus, NDF, life
history, African elephant, resource limitation, productivity.
vi
CHAPTER 1
GENERAL INTRODUCTION
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1.1 Introduction
One of the major concerns regarding elephant conservation and management is
the impact that they have on vegetation in enclosed areas. This issue can be
extended to the problem that elephants, both individually and as populations,
may become limited by resource availability. In accordance with optimal foraging
theory, elephants should consume the highest quality food sources first,
switching to lower quality foods as the abundance of the higher quality foods
decreased. This would result in them consuming food of declining quality over
the years, possibly to the point where sufficient nutrients to maintain condition
are no longer available, thereby resulting in a decrease in body condition. This
may then extend to reduced fitness, as predicted by density dependence. This
project addresses this hypothesis.
1.2 The elephant problem

The concept of the “elephant problem” was first introduced by Caughley (1976a),
referring to the trend of elephants transforming forest to open savannah or
grassland. The concern was that elephants were destroying trees faster than
they could regenerate, thereby altering habitats irreversibly. This concern was
based on the idea that ecosystems should be maintained at some balanced state,
and that any changes would negatively affect biodiversity. Although the idea of a
stable equilibrium in ecosystems has since been replaced with the view that
disturbances are essential to the maintenance and generation of biodiversity
(Connell 1978), these disturbances must still be assessed to ensure they comply
with the general primary objective of biodiversity conservation across a range of
organizational levels.
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Due to their large body size, elephant feeding and breakage impact on plants are
much more severe than smaller-sized herbivores (Owen-Smith 1988). These
impacts can also extend over larger areas than those by other herbivores, and
due to their longevity, can occur for much longer time periods (Laws 1970).
Although elephants have a relatively low specific metabolic rate and therefore
their relative daily dry weight food intake is low, their large size and low digestive
efficiency results in them requiring up to 180 kg of wet weight in food on a daily
basis (Owen-Smith 1988). In addition to this, elephants are known to be very
wasteful feeders, sometimes discarding the equivalent of between a quarter and
a half of the mass consumed (Paley 1997; Lessing 2007). They have been
recorded to fell and uproot trees with basal diameters of up to 60cm, only to feed
on the branch tips or roots, if at all (Chafota 2007).
Their tusks and trunks are well-adapted to feeding and allow them to access
plants and plant parts that would otherwise be unavailable to them. They use
their tusks to strip bark from trees, to dig into the trunks of baobabs (Adansonia
digitata) or to dig up roots (Barnes 1982; Lessing 2007). Their trunks allow them
to obtain very high intake rates (up to 2 kg.min-1 in thicket; Lessing 2007) by
enabling them to handle and chew food simultaneously. It also allows them,
combined with their shoulder height, to feed up to 8m above the ground (Croze
1974).
Elephants consume a varied diet which includes a range of plants and plant parts,
and their feeding is generally presumed to be an important mechanism in
structuring plant communities (Laws 1970; Stuart-Hill 1992; Trollope et al. 1998;
Lombard et al. 2001). This emphasises the importance of understanding
elephant diets and dietary preferences. Numerous incidences of elephant-
induced vegetative changes have been recorded in both savannah and thicket
ecosystems (Savidge 1968; Laws 1970; Penzhorn et al. 1974; Barratt & Hall-
Martin 1991; Barnes et al. 1994). They are also known to severely impact the
plant resource base, especially where they occur at higher densities (Laws 1970;
2
Barratt & Hall-Martin 1991). Their impact on plant communities are not only a
matter of food requirements, however, as these do not account for their wasteful
feeding or their indirect impact on non-browsed species through trampling and
path formation (Landman et al. 2008).
In addition to their effects on plant communities, elephants also impact individual

plant and animal species. Elephants are the only herbivores capable of killing
adult baobab trees and have been linked to the decline in their numbers
(Swanepoel 1993; Barnes et al. 1994). They also have been linked to the decline
of several Acacia species (Van Wyk & Fairrall 1969; Pellew 1983; Barnes 2001),
marula trees (Sclerocarya birrea) (Jacobs & Biggs 2002), Aloe africana
(Penzhorn et al. 1974), mistletoes (Magobiyane 2006) and geophytes and dwarf
succulent shrubs in thicket (Moolman & Cowling 1994). In addition, they
negatively impact a wide range of animal species including rhino (Kerley &
Landman 2006), ants, birds, bats (Cumming et al. 1997) and ungulates (Owen-
Smith 1988; Fritz et al. 2002).
The degree to which elephants impact their environment is a function of their

movement and density. Elephants do not use the landscape in a uniform fashion,
thereby varying their impacts across the landscape and creating heterogeneous
patterns of biodiversity (Owen-Smith et al. 2006). Movement, and thus impacts
across the landscape is determined by topographic relief (Wall et al. 2006), the
utilization of different vegetation types (Guldemond & Van Aarde 2007), as well
as the distribution of surface water (Laws 1970; Owen-Smith 1996). Higher
densities are also associated with more rapid rates of biodiversity loss and
habitat degradation (Laws 1970; Cumming et al. 1997).
Elephants are often confined to relatively small conservation areas. This further
amplifies their effect on the environment, as it does not allow for their impacts to
be distributed through space and time, thereby hampering vegetation recovery
(Novellie et al. 1991) and increasing the probability of encountering vulnerable
3
species (O’Connor et al. 2007). Additionally, the lack of space for range
expansion often results in the formation of larger family groups, creating very
high local densities with subsequent catastrophic effects on the resource base
(Laws 1970).
The nature of their diet, their large size and food requirements, feeding behaviour
and adaptations, as well as their movement patterns thus allow elephants to
influence various aspects of their environment in a unique way. Their impacts on
populations and species subsequently affect ecosystem patterns and processes,
such as nutrient cycling, soil resources, vegetation structure and dynamics
(Hatton & Smart 1984; Owen-Smith 1988; Kerley et al. 1999; Cowling & Kerley
2002; Skarpe et al. 2004; Lessing 2007), and ultimately influence biodiversity
(Van Wyk & Fairall 1969; Penzhorn et al. 1974; Barratt & Hall-Martin 1991;
Moolman & Cowling 1994; Lombard et al. 2001; Kerley & Landman 2006). This
demonstrates the vital importance of understanding elephant populations in order
to manage them.
1.3 Elephant life history and population regulation

Elephant are characteristic of long-lived (K-selected) species with a low intrinsic
rate of increase, low reproductive rates, low mortality and low rates of population
turnover (Laws 1981). The mean age of first reproduction in South Africa
averages 11.3 years, with females reaching sexual maturity from the age of
seven onwards (Van Aarde et al. 2007). Gestation lasts for 22 months, with a
mean intercalving interval of 3.6 years in South Africa (Van Aarde et al. 2007).
Females may remain reproductively active well into their 50’s (Whitehouse &
Hall-Martin 2000) with life expectancies of over 60 years (Van Aarde et al. 2007).
There is a high level of maternal investment and survival rates are generally high
among all age classes (Van Aarde et al. 2007).
Large mammal population regulation generally occurs via one of two

mechanisms- either through resource limitation (bottom-up) (Fowler 1987) or
4
predation (top-down) (Messier 1994). Predation works via the direct reduction of
population size through the killing of individuals. Due to their large size, however,
elephants have few predators, and it thus unlikely that predation plays an
important role in population regulation (Van Aarde et al. 2007). Resource
limitation, on the other hand, works via density dependence where, as population
size increases, the availability of resources declines (Klein 1968; Sinclair 1974;
Owen-Smith 2002). As resources become limiting, population growth is slowed
due to changes in vital rates. According to Fowler (1987) vital rates have different
sensitivities to changes in population density, with an increase in juvenile
mortality usually being the first response, followed by an increase in the age at
first reproduction, an increase in the intercalving interval and finally an increase
in the mortality rate of adults. This is supported by Eberhardt (2002), who found
similar sensitivities in vital rate responses. For large mammal populations, these
changes occur at densities close to the carrying capacity (Fowler 1981) and have
been demonstrated in numerous studies (Buss & Savidge 1966; Laws & Parker
1968; Laws 1969; Corfield 1973; Hanks & McIntosh 1973; Sinclair 1974; Clutton-
Brock et al. 1983). Such density dependent effects are also often exacerbated by
the heterogeneous distribution of resources through space and time (Chamaillé-
Jammes et al. 2008).
It has been suggested that, in open systems, elephant populations may respond
to increased density and food limitation primarily through dispersal, with changes
in reproduction and survival rates occurring more slowly (Chamaillé-Jammes et
al. 2008). However, reserve sizes in South Africa preclude the possibility of
dispersal as a mechanism of response. Additionally, although some evidence in
support of density dependence in Kruger has been found (Van Aarde et al. 1999),
Van Jaarsveld et al. (1999) found no evidence of density dependence on
population growth in 5 recovering populations in South Africa based on survival
rates. This is supported by Gough & Kerley (2006), who found no evidence of
density dependence at the population level in the Addo Elephant National Park
(AENP), even at densities far exceeding the suggested carrying capacity.
5
This raises the issue that elephants are able to impact and alter their
environments with seemingly no ill effect on their own current wellbeing and
survival, and highlights the need to investigate elephant populations more closely
in order to detect small, but critical responses to resource limitation. This is
especially true for populations in which other methods of control, i.e. culling,
contraception or translocation, are not possible.
1.4 Caughley’s framework

According to Caughley’s (Caughley 1976b) plant-herbivore framework, an
interferential model would best explain the interactions between elephants and
their environment (Fig. 1.1).
Interferential
Plant Elephant
Associated herbivores/
biodiversity
Fig. 1.1 A graphical representation of Caughley’s interferential plant-herbivore

framework.
It suggests that elephants, through their nature as large herbivores, would have
an impact on their plant resource base, and that this impact would also affect
other associated herbivores and biodiversity (as has been illustrated by
numerous cited studies above). Through increased density and impact on their
resources, there should then also be an effect of plants on elephants through
resource limitation. One such way in which interaction would become apparent is
through density dependent effects on population performance. As has been
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noted above, however, little evidence of such processes has as yet been
documented in elephant populations.
1.5 Body condition as indicator of response

Another way in which resource limitation could be detected is by monitoring
changes in the body condition of individuals within a population. As resource
availability and/or quality declines beyond the level of maintenance requirements
there should be an associated decline in the body condition of the individuals.
This is based on one of the characteristics of inanition, whereby an animal tends
to become thin as it drops in condition (Harris 1945; Riney 1960; Hanks 1981).
By monitoring body condition over time it may be possible to detect periods,
populations or segments of populations that are resource limited. This will allow
for the identification of opportunities when management interventions may most
effectively influence demographics.
1.6 Elephants as herbivores

Elephants are classified as large, generalist mixed feeders, consuming different
proportions of browse and graze according to their availability (Owen-Smith 1988;
Ulrey et al. 1997; Codron et al. 2006). They are hindgut fermenters, and as such
make use of a large caecum inhabited by microorganisms to digest plant fibre
(Ulrey et al. 1997). They have a rapid gut transit time (~24hrs; Clauss et al. 2007),
and dry matter (DM) digestibility in free-ranging elephants has been estimated to
be 30-45% (Meissner et al. 1990), with an adult daily dry matter intake
requirement of 1-1.5% of body weight (Ulrey et al. 1997). The natural diet is
characterized by a very high fibre content, with a neutral detergent fibre (NDF)
content of 50-70% DM, and a crude protein content of 6-12% DM (McCullagh
1969b; Meissner et al. 1990; Hatt & Clauss 2006), although calculations by Ulrey
et al. (1997) based on extrapolations from nutrient requirements of horses
estimated crude protein requirements of up to 14% for lactating females and
growing juveniles.
7
Due to their gut morphology they are able to ingest large amounts of fibre without
slowing the throughput (Janis 1976) and this, combined with their rapid gut transit
time and large size (and thus low specific metabolic rate), enables elephants to
persist on a very low-quality diet (Demment & Van Soest 1985; Owen-Smith
1988; Duncan et al. 1990). As generalists they feed on a large variety of plant
species. A study in the AENP found 143 species included in their diet, five times
more than coexisting ruminant browsers (Kerley & Landman 2006). This enables
them to affect more plant species than any other mammal herbivore in the
Eastern Cape thickets (Kerley & Landman 2006).
1.7 Measuring diet quality

Although various other methods for the determination of diet quality exist, faecal
analysis has proven to be the most practical for the study of wild herbivores as it
is non-invasive and samples are easy to collect (Leslie & Starkey 1985). Faecal
samples are also representative of the diet consumed within the last day or two
(taking into consideration gut-transit times), allowing it to be used to assess diet
over time (Rees 1982). Diet quality is generally related to the amount of nitrogen
(i.e. protein), fibre and minerals (e.g. phosphorus, calcium) present in forage, and
studies have shown that values obtained in faecal samples are correlated with
those found in the diet (Moir 1966; Leslie & Starkey 1987; Ulrey et al. 1997).
Faecal analysis is therefore an easy and effective way of determining the diet
quality of elephants.
1.8 The elephant problem in the Eastern Cape

Despite the fact that the Addo elephant population density has more than
doubled since the fencing of the park and resource availability is seriously
declining (Kerley & Landman 2006), the elephant population continues to grow at
a mean rate of 5.8% per annum (Gough & Kerley 2007). This is thought to be
due to the nutritious, evergreen, drought-resistant nature of the thicket vegetation
which dominates the Eastern Cape (Everard 1987; Seydack & Bigalke 1992;
8
Stuart-Hill & Aucamp 1993). It has been predicted that the Addo elephant
population will continue to increase at a high rate at the expense of biodiversity
until such a point at which they irreversibly damage the accumulated resource
base (Gough & Kerley 2007).
The Addo elephant population is the longest established and best studied in the
Eastern Cape, and as such provides a good benchmark for the management of
other, more newly established populations within the province. Although these
other populations are presently still limited in size, they have the ability to
increase in the future (Kerley 2006). Their occurrence at various densities and in
areas differing in productivity provides me with an excellent opportunity to
investigate the effects of seasonal and site-specific changes in resource quality
on the body condition of elephants in both energetically and non-energetically
constrained life stages.
1.9 Rationale, objectives and research approach

The use of body condition indices may be a powerful approach to detecting
dietary resource limitation in elephants. This is particularly relevant in a species
of such large body size and slow demographic responses to environmental
perturbations. These body condition responses may reflect life-history, seasonal
and site specific foraging and resource constraints. This project therefore sets
out to address the following objectives:
1. Determine whether energetically constrained elephant life stages (i.e.

weaning, lactation and loss of tooth function) are more vulnerable to
resource limitations compared to non-energetically constrained life stages
(i.e. sub-adults and non-lactating females) in order to identify critical
periods of elephant life history.
9
2. Determine how seasonal changes in the quality and availability of
resources affect elephant body condition in order to identify critical periods
of resource limitation.
3. Determine how site differences in productivity and elephant density affect

elephant body condition in order to identify populations that may be
resource limited.
The research approach was to use the well-studied Addo elephant population to
identify life stage variation in body condition, and to use this and other Eastern
Cape populations to assess seasonal and site specific body condition responses
of elephant.
10
CHAPTER 2
GENERAL DESCRIPTION OF THE STUDY SITES AND
POPULATIONS
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2.1 Background and elephants in the Eastern Cape
2.2.1 Historic populations and distribution
The presence of elephants in the broader Eastern Cape region is well
documented over the last few centuries, with archaeological discoveries
providing evidence of their occurrence within the province up to 1000 years
before present (Plug & Badenhorst 2001). These discoveries are supported by
more recent (during the last 100 years) evidence, consisting mostly of bone
material, some rock paintings and sightings, and the extensive hunting records of
the occurrence of elephants within the thicket and thicket-associated habitats in
the Eastern Cape (Boshoff et al. 2002a; Skead 2007).
Fig. 2.1 Map illustrating the historic distribution of elephants in the broader
Eastern Cape region (from Skead 2007).
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The first written records of elephants for most parts of the region date from the
late 1700’s when ivory became a valuable commodity in great demand in Europe,
causing elephant hunting to increase in popularity. From all the available records
it would appear that elephants were once widespread throughout the Eastern
Cape (Fig. 2.1) occurring in the Thicket, Forest, Grassland and Savanna biomes,
and on a patchy basis in the Fynbos and Nama-Karroo biomes (Hall-Martin 1992;
Skead 2007), and that numbers in the order of 100 000 animals is likely to have
been present before 1652 (Hall-Martin 1992).
Fig. 2.2 Map illustrating the three zones of density distribution of elephants in the
Eastern Cape region (from Boshoff et al. 2002a).
Most accounts of high elephant densities are from areas of thicket. Thicket was
also the dominant vegetation type, comprised of evergreen trees and shrubs high
in nutrients (Boshoff et al. 2002a). This was found to be especially true for the
coastal and mesic succulent thicket, and where these thickets formed mosaics
with forest and savanna vegetation- conditions that were prevalent along the
coastal belt. A reconstruction of relative elephant densities, status and habitat
12
types for the Eastern Cape was mapped by Boshoff et al. (2002a), indicating the
three likely zones of elephant occurrence in the broader Eastern Cape, with high
densities of resident populations occurring along the coastal belt, lower densities
of mostly migrant and nomadic populations occurring in the sub-coastal belt, and
very low densities or absence in the inland zone (Fig. 2.2).
2.2.2 Decline of the elephant

The near demise of the Eastern Cape elephants was indisputably caused by the
arrival of European settlers, as the San, the Khoi and the Bantu-speaking people
only utilized elephants at very low levels (Boshoff et al. 2002a). The first group of
European hunters arrived in Algoa Bay in 1702 in search of ivory, and by the mid
1700’s hunting activity had spread as far east as Pondoland (Boshoff et al.
2002a). By the late 1700’s and early 1800’s hunting had all but destroyed the
populations in the Peddie-King William’s Town area, and by the mid 1800’s
elephants were “rarely seen” in the Albany and Bathurst districts (Boshoff et al.
2002a). Ivory markets were established at various locations in the early to mid
1800’s, and large amounts of ivory were exported from Port Elizabeth (Hall-
Martin 1992).
By 1850 the Addo and Zuurberg populations had been significantly reduced, and
elephants were no longer found in the areas of the Great Kei and Great Fish
rivers (Boshoff et al. 2002a). The last of the elephants in the Bathurst district
were shot during the first decade of the 20th century, and the last known account
of elephants in western Pondoland was in 1860 (Boshoff et al. 2002a). The
remaining Addo elephants (about 130) were directly persecuted in 1919 when
Major P.J. Pretorius was hired by the government to eradicate the herd, as they
were in conflict with local farmers and human settlements (Hoffman 1993). He
proceeded to kill about 120 individuals between June 1919 and July 1920,
sparing only 16 individuals. By the time the Addo Elephant National Park (AENP)
was proclaimed in 1931, the Addo population had declined to 12 individuals
(Whitehouse & Hall-Martin 2000). Of these, 11 individuals went on to form the
13
founding population, representing the sum total of elephants in the Eastern Cape
at that time (Hall-Martin 1992; Whitehouse & Hall-Martin 2000).
2.2.3 Current populations and distribution

Since the establishment of the AENP the presence of elephants in the Eastern
Cape has changed significantly. The Addo population has flourished, with the
construction of elephant-proof Armstrong fencing in 1954 (Whitehouse 2000),
several expansions of the park and maintenance of perennial water sources
allowing the population to increase to its current number of 415 as at 30
November 2008 (K. Gough, pers. comm.). Additionally, with the development of
technology to translocate elephants over long distances, new populations were
established in the Eastern Cape. The first of these was in Shamwari Private
Game Reserve in 1992, followed by a small population in the Double Drift Game
Reserve in 1995, and two more populations in Kwandwe Private Game Reserve
and Bayethe Private Game Reserve in 2001 (Hall-Martin et al. 2002). Since then,
these populations have grown in size and more populations have been
established across the Eastern Cape, with population sizes ranging from 3 to 415
individuals (Fig. 2.2).
Fig. 2.2 The rate of establishment of elephant populations in the Eastern Cape
over the last 15 years (from Kerley 2006).
14
Six of these populations were used in the current study (Fig. 2.4).
Fig. 2.4 Map showing the locations of the six study sites (ArcGIS 9.2; map units:
decimal degrees; not projected).
2.2 Study sites and populations

2.2.1 Addo Elephant National Park- Main Camp
2.2.1.1 Site description, topography and geology
The Addo Main Camp (AMC) is located at 33°31’S, 25°24’E, approximately 60
km north east of Port Elizabeth, south of the R342 road from Paterson to
Kirkwood (Fig. 2.4). It spans an area of 103 km2, and forms part of the Addo
Elephant National Park (AENP). The topography of the AMC is characterized by
a series of valleys and gently undulating ridges, which rise from 71 m.a.s.l. to 354
m.a.s.l. (Paley & Kerley 1998), with the Zuurberg limestone plateau dominating
the eastern edge of the AMC (Barratt & Hall-Martin 1991).The dominant
substrate on which the AMC lies is sandstone and mudstone from the Uitenhage
15
Series, which is covered by a layer of red-brown granular clay loam soil rich in
humus (Archibald 1955). The soils formed are neutral, fine-grained and relatively
fertile (Hoffman 1989). The Zuurkop plateau is covered by grey calcrete and red-
brown aeolian sands (Toerien 1972; Barratt & Hall-Martin 1991). A number of
small, natural pans and waterholes are scattered throughout the park, although
these only hold water for short periods after rain. Permanent water is supplied by
means of three artificial water holes and two dams.
2.2.1.2 Climate
The AMC is located within the semi-arid region of South Africa, with a mean
annual rainfall of 396 mm recorded between 1961 and 1990 at the Citrus
Research Station, approximately 3 km south-west of the AMC (SA Weather
Service 2008). Rainfall occurs throughout the year and is mostly associated with
post-frontal events (Hoffman 1989), with peaks in late summer (February-March)
and spring (October-November) (Fig. 2.5). Prolonged droughts occur on a
regular basis (Barratt & Hall-Martin 1991).
50
45
40
35
Rainfall (mm)
30
25
20
15
10
5
0
J F M A M J J A S O N D
Months
Fig. 2.5 Average monthly rainfall recorded at the Citrus Research Station for the
period 1961-1990.
16
Mean maximum temperatures range from c. 29°C in the summer to c. 22°C in the
winter, although temperatures in excess of 40°C are often recorded in summer
(Stuart-Hill 1992).
Fig. 2.6 Map of the Addo Main Camp depicting the five main vegetation types
(ArcGIS 9.2; map units: decimal degrees; not projected, Mucina & Rutherford
2006). See Fig. 2.4 for location.
2.2.1.3 Vegetation
The AMC vegetation is classified as Xeric Succulent Thicket, which in its
undisturbed state is characterized by a dense tangle of succulent and spinescent
shrubs and lianas reaching 2-4 m (Paley & Kerley 1998; Henley 2001). Sundays
Thicket, dominated by Spekboom (Portulacaria afra), covers more than 66% of
the AMC (Fig. 2.6). Coega Bonteveld, consisting of a mosaic of low thicket with
secondary open grassland, Albany Coastal Belt, dominated by short grassland
with scattered bush clumps, and some Albany Alluvial Vegetation and Kowie
Thicket is also present (Mucina & Rutherford 2006). Dominant shrubs and low
trees include Azima tetracantha, Capparis sepiaria, Carissa haematocarpa,
Gymnosporia spp., Rhus spp., Euclea undulata and Schotia afra, with a high
17
incidence of Cynodon dactylon and Platythyra haeckeliana in the grassland
dominated areas (Landman et al. 2007).
2.2.1.4 Elephant population history

The establishment of the AENP in 1931 allowed for the allocation of 22.7 km2 for
the conservation of the 12 remaining elephants, although one was killed when
the animals were herded into the park area, leaving a founder population of 11
(Whitehouse & Hall-Martin 2000). This number increased very slowly to 22
elephants in 1954, at which time elephant-proof fencing was introduced
(Whitehouse & Hall-Martin 2000). After 1954 the population grew at an
exponential rate, reaching 284 individuals in 1998 (Whitehouse & Hall-Martin
2000), and currently numbering 415 (K. Gough, pers. comm.). During this time
there have been five expansions of the AMC, as well as the translocation of 4
Kruger bulls into the camp in 2002, and 63 individuals out to the Nyathi
Concession Area (NCA) in 2003, with corresponding changes in elephant density
(Gough & Kerley 2006). For most of the history of the park, the elephant density
has remained well above the 2 elephants/km2 estimated ecological carrying
capacity for subtropical thicket vegetation (Barratt & Hall-Martin 1991; Novellie et
al. 1996), as well as the 0.54 elephants/km2 calculated by Boshoff et al. (2002b).
The current density stands at 4.03 elephants/km2 (K.Gough, pers. comm.).
2.2.1.5 Elephant diet

The most recent elephant dietary study was conducted by Landman et al. (2007).
From this 13 Principal dietary items (PDI- species contributing 3% or more to the
diet) were identified, which made up 66.5% of the total diet. Of these, 7 species
contributed more than 3% each, namely Cynodon dactylon (19.6%), Portulacaria
afra (9.3%), Carissa haematocarpa (6.1%), Panicum deustum (4.5%), Azima
tetracantha (4.4%), Schotia afra (3.4%) and Eragrostis obtusa (3.2%).
18
2.2.2 Addo Elephant National Park- Nyathi Concession Area
The Nyathi Concession Area (NCA) is located to the north of the Addo Main
Camp and also forms part of the AENP (Fig. 2.4). It lies in the Coerney River
Valley, spanning an area of approximately 114 km2, and includes the southern
slopes of the Zuurberg Mountains (Davis 2005). For this reason, the terrain is
more rugged and mountainous than the AMC. The geology of the Zuurberg
Mountains is dominated by sandstones and quartzites of the Tafelberg and
Witteberg Series (Mucina & Rutherford 2006), with the south being dominated by
mudstones and shales of the Uitenhage group. Sandy soils dominate the
northern section of the park, with red loamy to clayey soils found throughout most
of the rest of the park (Mucina & Rutherford 2006). Water is supplied by the
Courney River, which runs in a south-westerly direction, as well as two artificial
dams and a couple of natural pans within the concession area.
2.2.2.2 Climate
The NCA also falls within the semi-arid region, with temperatures similar to the
Addo Main Camp, but is thought to receive more rainfall on average than the
AMC due to orographic effects (Davis 2005).
2.2.2.3 Vegetation
According to Mucina & Rutherford (2006) the vegetation of the NCA is dominated
by Sundays and Kowie Thicket, with Albany Alluvial Vegetation in the south. The
Zuurberg Mountain in the north is covered by a small section of Southern Mistbelt
Forest and Zuurberg Fynbos, characterized by grassy fynbos on the quartzite
dominated soils and closed ericoid shrubland/grassland on the shale dominated
soils (Fig. 2.7, Mucina & Rutherford 2006). The Sundays Thicket is
predominantly found on the low mountains and foothills, and is characteristically
tall and dense with many trees and shrubs (Mucina & Rutherford 2006).
Succulents and spinescent species, as well as a variety of lianas are common
(Mucina & Rutherford 2006). Kowie Thicket is found mainly on the drier, north-
19
facing slopes and is dominated by succulent euphorbias and aloes with a thick
understory of thorny shrubs, woody lianas and shrubby succulents (Mucina &
Rutherford 2006).
Important plants include Schotia afra, Cussonia spicata, Euclea undulata, Olea
europaea, Azima tetracantha, Portulacaria afra, various Viscum and asparagus
species, as well as numerous woody succulent and woody climbers (Mucina &
Rutherford 2006). Dominant grasses include Cynodon dactylon, Eragrostis
obtusa, Panicum maximum and P. deustum (Mucina & Rutherford 2006). A large
number of herbs, succulent herbs and geophytes are also present.
Fig. 2.7 Vegetation map of the Nyathi Concession Area showing the six main
vegetation types (ArcGIS 9.2; map units: decimal degrees; not projected, Mucina
& Rutherford 2006). See Fig. 2.4 for location.
20
The NCA was first stocked with elephants from the Addo Main Camp in May
2003, when 63 individuals were moved across the R342 Paterson to Kirkwood
road and fenced in (K. Gough, pers. comm.). The group consisted of 17 mature
females, nine sub-adult females, one sub-adult male, 20 calves, and 15 mature
bulls. An additional four males were introduced into the park from Kruger in 2003,
but one subsequently died in 2004. At the time of stocking the density was 0.45
elephants/km2. Before this, elephants had been absent from the NCA for more
than 50 years (Davis 2005). Since then the population has grown to 88 (K.
Gough, pers. comm.) and a density of 0.77 elephants/km2.

According to Davis (2005), eight plant species were found to make up the
principal dietary items (PDI) of the elephants in the NCA, and included Azima
tetracantha, Cynodon dactylon, Panicum deustum, Cussonia spicata, Viscum sp.,
Rhiocissus digitata, Schotia afra and a category made up herbaceous plants
which could not be identified to the genus level.
2.2.3 Asante Sana Private Game Reserve

Asante Sana Private Game Reserve (ASPGR) is located at 32° 18’ S, 24° 58’ E,
approximately 40 km east of Graaff-Reinet in the basin of the Sneeuberg
Mountains (Fig. 2.4). It spans an area of 108 km2, and includes mountains
reaching up to 2 100 m.a.s.l. with slopes of varying steepness, as well as the
basin floor, lying at approximately 1000 m.a.s.l. (Boshoff & Kerley 1997). The
geology consists of the Beaufort Group of the Karroo Sequence, with greenish
grey and red mudstones and sandstones of the Middleton formation overlain by
greenish grey and red mudstones and shales and sandstone of the Balfour
formation, with dolerite sills of varying thickness intruding the Balfour Formation
(Boshoff & Kerley 1997). The basin floor consists mainly of alluvial soils, which
are thought to be important from an agricultural point of view (Hill 1993 as cited
21
by Boshoff & Kerley 1997). These are dominated by loamy sand to loamy duplex
type soils, with high amounts of potassium, copper and boron, and low levels of
zinc (Boshoff & Kerley 1997).
2.2.3.2 Climate
ASPGR occurs in a region where most rainfall is received in the summer months
(October – March) (Venter et al. 1986). The mean annual rainfall recorded for the
closest town, Graaff Reinet was 361 mm for the period 1961-1990 (Fig. 2.8, SA
Weather Service 2008). It is thought that the rainfall on the higher slopes of the
reserve should fall within the 600-700+ mm range (Boshoff & Kerley 1997).
Extended periods of drought are often experienced in this region, however, this is
partially alleviated by infrequent winter rains and run-off from melting snow
(Boshoff & Kerley 1997).
70
60
50
Rainfall (mm)
40
30
20
10
0
Month
Fig. 2.8 Average monthly rainfall recorded at Graaff Reinet for the period 1961-
1990.
The mean annual temperature ranges between 15 and 17.5°C, although

temperature extremes are experienced both in summer (< 30°C) and winter
(>0°C) (Venter et al. 1986). Frost is occasionally recorded (mean annual
frequency of 5 days) on the basin floor and valley bottoms (Boshoff & Kerley
1997).
22
2.2.3.3 Vegetation
According to Mucina & Rutherford (2006) two vegetation types can be discerned
in Asante Sana, namely Camdeboo Escarpment Thicket and Karoo Escarpment
Grassland (Fig. 2.9).
Fig. 2.9 Vegetation map of Asante Sana illustrating the two main habitat types
Merxmuellera disticha and Themeda triandra dominate the grassland, whilst the
Escarpment Thicket is dominated by Acacia karroo, Celtis africana and Grewia
robusta (Mucina & Rutherford 2006). Pentzia incana, Pentzia spinescence and
Eriocephalus ericoides dominate the Karroid Dwarf Shrubland, whilst the
dominant species in the Riverine Thicket include Acacia karroo, Rhus lancea and
Cynodon dactylon (Boshoff & Kerley 1997), with Portulacaria afra, a
characteristic of this vegetation type, only occurring sparsely.

After an initial study by Boshoff & Kerley (1997) to investigate the suitability of the
area to elephants, a small family group of nine elephants was translocated to the
23
area from Sabi-Sands Private Game Reserve in 2004. The group consisted of
three mature females, four sub-adult females, as well as two calves (both about
two years old). Shortly after translocation one of the sub-adult females as well as
a male calf died. A calf was born on the reserve in June 2004, followed by
another calf in January 2005 to bring the population back up to nine individuals.
Due to the lack of a sexually mature bull in the group and the unchanged reserve
size, the population size and density, has remained constant at 0.083
elephants/km2.

According to a dietary study by Minnie (2006), 66 species make up the elephant
diet in the ASPGR, with 17 species being identified as PDI. These PDI were
dominated by three species, namely Acacia karroo (24.7%), Cynodon dactylon
(8.5%) and Grewia robusta (8.3%), contributing 42.7% to the total diet.
2.2.4 Shamwari Private Game Reserve

Shamwari is located at 33°20’S, 26°01’E, along the N2 national road to Port
Elizabeth, approximately 11 km east of Paterson (Fig. 2.4). It spans an area of
approximately 200 km2, of which 180 km2 is available to elephants. It is
characterized by a series of undulating hills ranging from 196 m.a.s.l. in the south
to 628 m.a.s.l. in the north. According to O’Brien (2004), the main geological
formations found in Shamwari include Bokkeveld Series Shale, Witteberg
Quartzites, Karroo Sandstone and Sundays River Formations. Because of the
steep elevation gradient four main substrate types are exposed and available to
plant communities, namely shale, sandstone, quartzite and calcrete. Some
deeper alluvial soils also occur on the lower lying land (O’Brien 2004). Water is
supplied by the semi-perennial Bushman’s River, which flows through the
reserve, as well as numerous other pans and dams scattered throughout the
reserve.
24
2.2.4.2 Climate
Shamwari falls between the summer and winter rainfall areas of the Eastern
Cape and thus receives rainfall throughout the year (Fig. 2.10), with peaks in
March (63.3 mm), August (63.5) and November (58 mm). Annual precipitation for
the period 1999 to 2007 averaged 550 mm (Shamwari rainfall records).
70
60
50
Rainfall (mm)
40
30
20
10
0
Month
Fig. 2.10 Average monthly rainfall recorded at Shamwari for the period 1999-
2007.
Temperature data for Shamwari, or the nearest town, Paterson, were not
available, and therefore temperature data from Addo (approximately 40 km
south-west of Shamwari) were used. As indicated above (AMC section) the mean
maximum temperatures ranged from approximately 29°C in summer to
approximately 22°C in winter, although temperatures in excess of 40°C are often
recorded in summer (Stuart-Hill 1992).
2.2.4.3 Vegetation
Shamwari encompasses 5 vegetation types, including Kowie Thicket, Bisho
Thornveld, Suurberg Quartzite Fynbos, Albany Coastal Belt and Suurberg Shale
Fynbos, with Kowie Thicket being the dominant vegetation type (Fig. 2.11;
Mucina & Rutherford 2006). A study done by O’Brien (2004) identified a further 8
vegetation types, including Grassland, Bushclump Savanna and Riverine Thicket.
25
Fig. 2.11 Vegetation map of Shamwari depicting the five major vegetation types
According to O’Brien (2004) the thicket is characterized by a dense tangle of

evergreen succulent and woody shrubs and trees dominated by species such as
Schotia afra, Carissa bispinosa, Cussonia spicata, Pappea capensis, Euclea
undulata, Aloe africana, A. ferox, and Portulacaria afra. The grasslands are
dominated by Themeda triandra, Heteropogon contortus and Eragrostis curvula,
while species such as Pennisetum clandestinum, Cenchrus ciliarus and Cynodon
dactylon can be found on the old farm lands. The composition of woody species
is similar in the bushclump savanna as in the subtropical thicket, with
bushclumps being interspersed with grasses such as Digitaria eriantha, Setaria
neglecta, Eustachys paspaloides, Themeda triandra and Eragrostis curvula
26
(O’Brien 2004). The Albany coastal belt includes species such as Erythrina caffra,
Euphorbia triangularis, Grewia occidentalis and Rhus lucida, whilst the fynbos
areas support ericoid shrubland and grassland species (Mucina & Rutherford
2006).

According to O’Brien (2002) the first elephants were introduced between 1992
and 1993, when 14 orphans were translocated from the Kruger National Park,
consisting of seven males and seven females. An additional group of eight
orphans were introduced from Mpongo Park in East London in 1994, comprising
four males and four females. In 1997 a family herd of nine elephants were
reintroduced, with only seven surviving (two males and five females). Two
females were subsequently brought from Knysna in 1999. In 2003, 13 individuals
were relocated to Bushman Sands Game Reserve and Sanbona Wildlife Centre,
and two adult bulls were euthanased. Between 2003 and 2008 another 25
individuals were translocated out of the park, six to Sanbona Wildlife Centre, five
to Mpongo Parks, three to Kamala Game Reserve, six to Bushman Sands Game
Reserve and five to Rippons Game Reserve. The current population stands at 47
individuals and a density of 0.26 elephants/km2.

According to Roux (2006) the elephant diet in Shamwari consists of 23 species
from 19 families, with the bulk of the diet made up of Acacia karroo (36%), Rhus
spp. (11%), Opuntia ficus-indica (10%), Azima tetracantha (9%), Gymnosporia
spp. (6%), Pappea capensis (3%), Schotia afra (3%) and Combretum caffrum
(3%).
2.2.5 Kariega Private Game Reserve

Kariega is located in the Kariega River Valley (Fig. 2.4), along the R343 regional
road, approximately 45 km south of Grahamstown (33°35’S, 26°37’E). It spans
27
an area of 190 km2 and ranges from an altitude of 23 m.a.s.l. at the base of the
Kariega River Valley to 262 m.a.s.l. at the northwest corner of the reserve. The
northern half of the reserve is located on a plateau situated above the Kariega
River Valley, with the southern half consisting of undulating hills in the west and
flat, low-lying ground in the east. The principal geological formations of the
reserve include Beaufort Group shale, mudstone, solonetic soils and sandstone,
and Cape Supergroup sandy clays and lithosoils (Low & Rebelo 1996). Apart
from several small dams, water in the park is mainly supplied by the perennial
Kariega River, which runs through the park for 11 km.
2.2.5.2 Climate
Kariega falls within the spring-dominated rainfall area of the Eastern Cape, but
has a pronounced bi-modal rainfall pattern (March-April and November-
December) (Stone et al. 1998). The average annual precipitation for the closest
town for which data were available, Port Alfred, for the period 1936 to 2003 was
640 mm (Fig. 2.12; SA Weather Service 2008). Its close proximity to the coast
results in Kariega having a slightly higher average rainfall than other reserves in
the Eastern Cape due to coastal fog, where moist air from the sea moves over
the cold land surface (Stone et al. 1998).
Kariega’s close proximity to the coast also results in it being affected by land and
sea breezes in the late afternoons/evenings, caused by differential heating and
cooling of the land and sea (Stone et al.1998). These breezes moderate the
temperatures by decreasing day time temperatures and increasing night time
temperatures, thereby also decreasing the amount of frost exposure in the
reserve. Temperature data were not available for Kariega, or the closest town,
Bushman River’s Mouth, and therefore data from Port Alfred were used. The
mean maximum temperature for summer was approximately 27°C and
approximately 20°C for winter, with temperatures in excess of 30°C and below
5°C occasionally recorded (SA Weather Service 2008).
28
80
70
60
Rainfall (mm)
50
40
30
20
10
0
Month
Fig. 2.12 Average monthly rainfall recorded at Port Alfred for the period 1936-
2003.
2.2.5.3 Vegetation
Three major vegetation types are present on the reserve (Fig. 2.13), namely
Kowie Thicket, Albany Coastal Belt and Cape Estuarine Salt Marshes (Mucina &
Rutherford 2006), with Kowie thicket being the dominant vegetation type,
occurring on most of the slopes in the reserve (Parker 2004).
Characteristic species include Cassine aethiopica, Euphorbia triangularis, E.

tetragona and Plumbego auriculata (Low & Rebelo 1996). Acacia cyclops has
also invaded some of this vegetation type on the reserve. Some old farm land
occurs on the flood plains of the Kariega River, and is characterized by grass
species such as Panicum stapfianum, Eragrostis curvula and Themeda triandra,
with some planted exotic trees also present (Parker 2004). Albany coastal belt
occurs along the eastern and western boundaries of the reserve in deep valleys,
and includes species such as Mimusops caffra, Apodytes dimidiata, Sideroxylon
inerme and Cassine aethiopica (Low & Rebelo 1996). Cape salt marshes make
up only a small fraction of the reserves southern boundary.
29
Fig. 2.13 Vegetation of Kariega showing the three main vegetation types (ArcGIS
9.2; map units: decimal degrees; not projected, Mucina & Rutherford 2006). See
Fig. 2.4 for location.

Elephants were introduced to Kariega in 2004 when a family group of 11
individuals were translocated from the Sabi-Sands Private Game Reserve. The
group consisted of seven females and four males, of which three were adults, six
were sub-adults, and two were 5 year-old calves. Additionally, two adult bulls
were introduced in 2005 from the AENP, and six calves have since been born on
the reserve (one each in 2005, 2006, and 2007, and three in 2008). In early 2008
one of the adult bulls died from a tooth abscess, leaving the current population of
19 elephants at a density of 0.1 elephants/km2.

Wolmarans (2006) identified 15 PDI’s in the elephant diet. These included
Cynodon dactylon (7.8%), Rhus crenata (5.6%), Azima tetracantha (5.3%) and
Schotia afra (5.3%) as the dominant species. With the exception of the high
occurrence of C. dactylon, woody shrubs dominated the diet in all seasons.
30
2.2.6 Blaauwbosch Private Game Reserve
Blaauwbosch is located at 33°21’S, 24°49’E, approximately 5 km SW of
Wolwefontein on the R75 road from Port Elizabeth to Graaff-Reinet (Fig. 2.4). It
spans an area of 45 km2 with mountainous terrain dominating the northern and
eastern sections with slopes of varying steepness, with the south being
characterized by a typical Karroo basin. Altitude ranges from 854 m.a.s.l. in the
north east to 420 m.a.s.l. in the south. The geology consists of shales and quartz
sandstones of the Witteberg group in the northern and central regions of the park,
with the southern half of the park dominated by conglomerates and sandstones
of the Uitenhage group (Mucina & Rutherford 2006). The soils are predominantly
red, acidic, fine-grained clay soils with a high phosphorus and low nitrogen
content (Hoffman 1989). Water in the park is supplied by a number of ground and
cement dams fairly evenly distributed across the park.
50
45
40
35
Rainfall (mm)
30
25
20
15
10
5
0
Month
Fig. 2.14 Average monthly rainfall recorded at Pinelands Farm (33°19’S, 24°51’E)
for the period 1951-2006.
2.2.6.2 Climate
Blaauwbosch falls within the semi-arid region of South Africa and receives an
average of 300 mm rain per annum (Fig. 2.14, calculated from rainfall records
from a neighbouring farm spanning from 1951 to 2006). Although rain is recorded
31
throughout the year, there is a distinct bimodal pattern (Stone 1998), with peaks
in March (44 mm) and November (33 mm). Extended periods of drought are not
uncommon.
As no temperature data were available for the park or the closest town, average
temperatures were calculated from 29-year temperature data for Jansenville,
approximately 50 km north east of Blaauwbosch, and Uitenhage, approximately
100 km south east. Average maximum temperatures for summer and winter were
31.2°C and 21.3°C respectively, with minimum temperatures recorded at 17.1°C
and 6.3°C (SA Weather Service 2008).
Fig. 2.15 Vegetation map of Blaauwbosch Private Game Reserve illustrating the
three dominant vegetation types (ArcGIS 9.2; map units: decimal degrees; not
projected, Mucina & Rutherford 2006). See Fig. 2.4 for location.
2.2.6.3 Vegetation
Three major vegetation types are present in Blaauwbosch (Fig. 2.15), namely
Lower Karroo Gwarrieveld in the north, Groot Thicket in the central regions and
Sundays Thicket in the south. The Gwarrieveld vegetation is predominantly
32
shrubland with a sparse canopy of Euclea undulata, Boscia oleoides and Schotia
afra with isolated outcrops of Portulacaria afra, whereas the Groot Thicket is
found on moderate to steep slopes of the mountain, dominated by fairly dense
and closed low succulent thicket with a poorly developed grass component
(Mucina & Rutherford 2006). Species include Euclea undulata, Grewia robusta,
Carissa bispinosa, Portulacaria afra, Crassula ovata and Viscum spp.. The
Sundays Thicket is as described in section 2.2.1.3 and 2.2.2.2.

A family group of eight elephants were translocated from the Sabi-Sands Private
Game Reserve to Blaauwbosch in 2005. The group was comprised of four males
and four females, of which three were adults, two were sub-adults and three
were calves. Two calves were subsequently born in 2006, bringing the total
population number up to 10 individuals. With the absence of an adult bull for
breeding, the density has since remained stable at 0.22 elephants/km2.

No elephant dietary study has yet been conducted in Blaauwbosch. Therefore
the observations of the reserve manager and field rangers were used as an
indication of the species most utilized by elephants. These included Acacia
karroo, Azima tetracantha, Opuntia ficus-indica, Euclea undulata and Pappea
capensis.
33
CHAPTER 3
LIFE-HISTORY VARIATION IN BODY CONDITION
¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯
3.1 Introduction
3.1.1 Body condition
The condition of an animal can be defined as a measure of the variation of its
body fat reserves and is essentially a reflection of its energetic state (Pitt et al.
2006). A minimum amount of energy is needed for maintenance, with any surplus
energy being allocated to growth, reproduction and storage (fat deposits). An
individual with a larger energy reserve will thus have more energy to allocate to
reproduction and fat storage, thereby increasing its chances of survival and
reproduction, and will thus ultimately experience increased fitness (Klein 1968;
Sinclair & Duncan 1972; Miller & Hickling 1990; Choquenot 1991; Hodges et al.
1998). Factors that may affect the condition of an individual can broadly be
classed into two categories, namely intrinsic and extrinsic. Intrinsic factors
include factors that are an inherent part of the animal, i.e. life-history stages (i.e.
pregnancy, lactation, weaning & loss of tooth function) and genetics, whilst
extrinsic variables refer to those factors independent of the animal, i.e. population
density, stress, forage quality production and rainfall (wet/drought periods).
3.1.2 Body condition over lifetime

During its lifetime an individual may be subjected to many changes in extrinsic
(ecological) variables. These include periods of changeable rainfall (drought/wet
periods) with associated changes in vegetation quality (Phillipson 1975; Malpas
1977; Grant et al. 1995). It may also be subjected to periods of stress (predation,
disease, human conflict), and variations in population density with associated
effects on food availability (Klein 1968; Skogland 1985; Choquenot 1991; Hodges
et al. 1999; Gough & Kerley 2006). Depending on the duration and severity,
these periods may impact the body condition of the individual through a decrease
in the quality and quantity of available food (Phillipson 1975; Grant et al. 1995).
34
An individual is also likely to experience periods of increased nutritional
requirements during certain life stages. These include weaning, when the
dependent calf is slowly weaned from its mother and is forced to start foraging for
itself whilst still maintaining a high growth rate (Lochmillar et al. 1982; Robbins
1983; Hayssen 1993), lactation, when the adult female bears the cost of the
nutrient-rich milk for her calf (Young 1976; Malpas 1977; Robbins 1983; Clutton-
Brock et al. 1989; Hayssen 1993; Reuter & Adcock 1998) and old age, when the
tooth surface starts wearing down, hampering forage mastication and thus
nutrient absorption (Laws 1981; Price & White 1985; Skinner & Chimimba 2005).
If there is not sufficient quality and quantity of forage available during these times
to meet the increased nutritional requirements, the animal will be forced to
mobilize body fat reserves and metabolize muscle tissue (Price & White 1985).
This will cause a decline in body condition and make the individual increasingly
more vulnerable to environmental stressors (McGregor & Butler 2008). This is
known as the “multiplier effect”, whereby a small deficiency in resources can
become lethal through the intercession of predators and pathogens (Owen-Smith
2002). An example of this increased vulnerability at these life stages can be
found in the Tsavo elephant die-off of 1970/71, where neonates, mothers with
calves and old individuals were the first to succumb to starvation during the
drought (Corfield 1973).
3.1.3 Condition measures

Various indices have been developed to measure the condition of an animal. The
majority of these use the amount of fat in various deposits throughout the body
as an index of body condition. This is based on the assumption that the quantity
of fat in a specific area of deposition is proportional to the total body fat reserves
in some predictive way (Sharp 1982). These indices include measures such as
the Kidney Fat Index (Malpas 1977; Choquenot 1991; Gallivan et al. 1995) and
Bone Marrow Fat (Malpas 1977). A number of other indices have also been used
to assess condition, namely blood chemistry and haematology (Malpas 1977;
Gallivan et al. 1995), adrenal cortical hypertrophy (Anderson et al. 1971), urinary
35
excretion of hydroxyproline (McCullagh 1969a; Malpas 1977), bioelectrical
impedance (Pitt et al. 2006) and body growth (Hodges et al. 1999). Many of
these methods are, however, invasive or impractical for studies on large
mammals. This has led to the development of visual body condition indices for
some mammals (Riney 1960; Albl 1971; Poole 1989; Grant et al. 1995; Reuter &
Adcock 1998; Wemmer et al. 2006), which have been found to give a reliable
reflection of overall condition. Additionally, visual condition estimations do not
only consider deposited fat reserves, but are partially dependent on changes in
muscle volume, thereby incorporating the deposition and mobilization of protein
resources (Sharp 1982). This is important, as animals losing condition will
catabolize muscle as well as fat reserves to meet their nutritional demands, even
in the early stages of weight loss (Price & White 1985; Owen-Smith 2002).
3.2 Hypotheses and aims

By comparing the body condition of energetically constrained life stages to non-
energetically constrained life stages (i.e. sub-adults and non-lactating females), I
am able to test the hypothesis that elephant body condition will vary as a function
of life stage, which can be visually represented (Fig. 3.1). Specifically, I predict
that individuals that are being weaned, lactating, or experience loss of tooth
function will have lower body conditions than those that are not energetically
constrained.
This will enable me to determine whether energetically constrained elephant life

stages (i.e. weaning, lactation and loss of tooth function) are more vulnerable to
resource limitations compared to non-energetically constrained life stages (i.e.
sub-adults and non-lactating females) in order to identify critical periods of
elephant life history.
Additionally, by comparing the body condition of individuals (within their

respective life stages) across seasons, I am able to test the hypothesis that
elephant body condition will vary seasonally. Specifically, I predict that there will
36
be a decrease in condition in seasons when resources may be limiting, and that
this seasonal variation will be reinforced by the various life stages.
Fig. 3.1 Expected changes in the body condition of a female elephant over her
lifetime.
3.3 Materials and methods

3.3.1 Selection of life-history groups
Only the AMC population was used to investigate the effect of individual life
stage on body condition. This was the only population for which all individuals
and their life histories are known (Whitehouse & Hall-Martin 2000; Gough &
Kerley 2006), and was also the only population large enough to allow statistical
analyses on the various life stages. A list of all the elephants was obtained (K.
Gough, unpubl. data) and these were sorted into the four categories investigated,
i.e. weaning, lactation, loss of tooth function, and sub-adults and non-lactating
females (pooled). Individuals were classified as weaning if they were under 4
years and had a younger sibling no older than 1 year, as the initial period of
weaning is thought to be the most energetically stressful. Although suckling up to
the age of 8 years has been recorded (Lee & Moss 1986), only adult females
with calves under the age of 3 years were classed as lactating, as the
37
intercalving interval in Addo averages 3.3 years (Gough & Kerley 2006). Adult
females older than 50 years were classified as experiencing loss of tooth function,
as the last set of molars erupt at about 47 years of age (Laws 1966). Sub-adults
were classified as males and females between the ages of 6 and 10 years (mean
age of first calving for the park calculated at 12.2 years (SD = 1.73) by Gough &
Kerley (2006)), with known-pregnant females under 10 excluded from this
category. Non-lactating females were classified as those females between the
ages of 10 and 50 that had not had a calf within the last 5 years. As not all
elephants meeting the criteria for each category were seen each season, sample
sizes differed between seasons. These are given in Appendix 1.
3.3.2 Data collection

Individual elephants were identified by ear notches and tusk shape, as well as
the unique wrinkle patterns on their faces and blood vessels in their ears
(Whitehouse & Hall-Martin 2000).This ensured that no individual was scored
twice in any given season, and that the changes in condition of the same
individuals could be tracked across seasons. Body condition scores were
obtained by visual observations of individuals within each population seasonally.
Body condition was scored as per Gough (in prep.), who expanded the visual
condition scale developed by Poole (1989), with an individual receiving a body
condition score of between 1 and 8, depending on the state of specific external
areas as follows:
(1) Emaciated: The shoulder blades, pelvic bone, spine and ribs protrude
heavily.
(2) Very thin: The shoulder blades, pelvic bone, spine and ribs protrude.
(3) Thin: The shoulder blades, pelvic bone, spine and ribs are noticeable.
(4) Fair: The shoulder blades, pelvic bone and spine are noticeable (the ribs
are not).
(5) Good: Slight sinking in front of and behind the pelvic bone is noticeable,
and the shoulder blades and spine protrude slightly.
38
(6) Very good: Slight sinking in front of and behind the pelvic bone is
noticeable, but the shoulder blades and spine is not (shoulders well
rounded).
(7) Fat: There is no sign of shoulder blades, pelvic bone or spine.
(8) Very fat: There is no sign of shoulder blades, pelvic bone or spine and fat
hangs from the belly.
Where possible, photographs were taken to aid in identification of individuals, as

well as to ensure consistency in the scoring of body condition over the study
period (see Fig 3.2 for example). Additionally, only one observer allocated the
condition scores over the study period, thereby reducing observer bias.
Fig. 3.2 An example of an elephant with a body condition score of 3 (photo top
right: note the visibility of the pelvic bone and ribs), and an elephant with a body
condition score of 7 (photo bottom left: there is no sign of the pelvic bone,
backbone or shoulder blades).
39
3.3.3 Data analysis
A three-way Log-linear analysis (Statistica 8.0; StatSoft Inc. 2007) was used to
test whether there was a significant difference in body condition between life
stages for the AMC, both within and between seasons. Factors were season, life
stage and body condition category. Significance was assessed at the 95%
probability level (Zar 1999).
Data were then explored graphically using Box-and-Whiskers plots (Statistica 8.0;
Statsoft Inc. 2007), with whiskers representing the 90% confidence interval (CI).
Where whiskers did not overlap, the comparison was taken to be statistically
significant. The 90% CI value was chosen, as the standard 95% CI would be too
conservative, thereby significantly increasing the probability of a Type I error
(Payton et al. 2003; Schenker & Gentleman 2001). According to Payton et al.
(2003) my chosen CI value would still be slightly conservative in detecting
significant differences.
Due to sample size limitations, the initial eight point body condition scale was
reduced to a four point scale (body condition ranking 3-6, for which most
observations occurred). This was done by eliminating rank 1, for which no
observations were recorded in any sample, and incorporating observations of
rank 2 into rank 3, and those of rank 7 and 8 into rank 6. It is felt that this
adjustment of the body condition scale would not significantly alter the pattern or
significance of the data, as the number of observations in ranks 2, 7 and 8 were
very small.
3.4 Results
The log-linear analysis revealed that a two-way model was adequate to explain
the data (χ2 = 114.93, df = 27, p < 0.001). The two-way interactions that were
significant were (1) season x body condition category, and (2) life stage x body
condition category. Thus, controlling for life stage, body condition was found to
differ significantly between seasons (χ2 = 57.63, df = 9, p < 0.001). Body
40
condition was also found to differ significantly between life stages, controlling for
season (χ2 = 51.79, df = 9, p < 0.001). See Appendix 2 for full results.
Graphical exploration of these results revealed that, in winter, lactating

individuals were found to have significantly lower body conditions than weaned or
non-energy stressed individuals (Fig. 3.3). Old individuals were also in poorer
condition in winter, but this was not significant due to the high variance in their
score for this category. Old individuals had significantly lower body condition
scores than weaned and non-energy stressed individuals in spring, with lactating
females having intermediate body condition scores as compared to the other life
stages (Fig. 3.4). In summer both old and lactating individuals had significantly
lower body conditions than weaned or non-energy stressed individuals (Fig. 3.5).
This trend was repeated again in autumn, with both old and lactating individuals
in poorer condition than weaned and non-energy stressed individuals (Fig. 3.6).
6.0
5.5
5.0
Body Condition
4.5
4.0
3.5
3.0
WOld WLact WWean Wsub/non
Fig. 3.3 Mean elephant body condition, ± 0.9 confidence interval, for the four life
stages for Addo in winter. First letter of x-axis label denotes season; last letters
denote life stage (i.e. old, lactating, weaned and sub-adult/non-lactating)
41
6.0
5.5
5.0
Body Condition
4.5
4.0
3.5
3.0
SpOld SpLact SpWean Spsub/non
stages for Addo in spring.
6.0
5.5
5.0
Body Condition
4.5
4.0
3.5
3.0
SuOld SuLact SuWean Susub/non
stages for Addo in summer.
42
6.0
5.5
5.0
Body Condition
4.5
4.0
3.5
3.0
AuOld AuLact AuWean Ausub/non
stages for Addo in autumn.
Body conditions were found to be the highest in winter for all life stages (Fig. 3.7).
After the initial decline in condition from winter to spring, all but the lactating
females showed an improvement in condition in summer. This was followed
again by a decrease in condition for all life stages, except lactating females,
which showed a slight increase in condition from summer to autumn.
43
6.5
6.0
5.5
Body Condition
5.0
4.5
4.0
3.5
3.0
2.5
SpOld
SuOld
AuOld
WLact
WOld
WWean
Wsub/non
SpWean
SuWean
AuWean
SpLact
Spsub/non
SuLact
Susub/non
AuLact
Ausub/non
Fig. 3.7 Mean body conditions, ± 0.9 confidence interval, for all life stages in the
AMC. Vertical lines separate the four different seasons, i.e. winter, spring,
summer and autumn.
The condition of old individuals did not show any significant seasonal variation,
although there was a general trend of decrease from winter to autumn. Lactating
individuals, however, had significantly lower conditions in summer and autumn
than in winter, with intermediate condition in spring. Weaned individuals were in
significantly better condition in winter than in spring and autumn, with summer
condition almost significantly lower than winter. Non-energy stressed individuals
(sub-adults and non-lactating females) were in significantly better condition in
winter than in any of the other seasons, with autumn condition being significantly
lower than summer and winter.
Within the old individuals (>50 years), a comparison between age and body
condition revealed a general decline in body condition with increasing age, with a
sudden decrease in condition from 57 years (Fig. 3.8). This relationship, however,
was not significant (p = 0.3695).
44
4.2
4.0
3.8
3.6
Body Condition
3.4
3.2
3.0
2.8
2.6
51 52 53 54 55 56 57 58 59 60
Age (years)
Fig. 3.8 The relationship between age and body condition for old individuals.
Points on the graph represent the average body condition score of each
individual for all seasons combined.
3.5 Discussion
Visual body condition estimates are not as widely used as other biological
measures of condition, such as Kidney Fat Index and Bone Marrow Fat. As they
are not measured by some hard-and-fast value, but rather by the observation of
the researcher, they are at risk of proving unreliable or inconsistent results. My
results, however, suggest that my visual determination of elephant body
condition has provided me with a reliable, repeatable, measure of condition,
giving an accurate reflection of the nutritional status of the individuals.
3.5.1 Condition and life stage

Although the non-energy stressed class (range: 4.4 to 5.7) exhibited better
conditions than the energy stressed classes (range: 3.3 to 5.6), this difference
was not significant for all energy stressed classes in all seasons. Weaned
individuals had almost identical condition estimates to the non-energy stressed
individuals in all seasons, ranging in condition from 4.4 to 5.6. This similarity in
45
condition suggests that there is no additional energy stress on these weaned
calves. This could be due to the extended period of mixed-feeding during
weaning observed in the African elephant (Langer 2003). This extended period of
milk supplementation may allow calves the time to adjust to obtaining sufficient
quantity and quality of solid foods to meet their own nutritional needs (Langer
2003). Thus, when their suckling efforts are increasingly rejected, or stopped by
the arrival of a new sibling, they are able to rely solely on solid food without
negatively affecting their condition. Another possible explanation is the tendency
for some elephant mothers to allow an older sibling to continue suckling after the
arrival of a new calf (Moss & Lee 2004). In such instances the older calf would
still receive some supplemental milk, providing some of the energy needed for
growth and the maintenance of condition.
Lactating females exhibited body conditions of between 4.0 and 4.8, and had
significantly poorer condition than non-energy stressed individuals in all seasons
except spring, when body condition was found to be lower, but not significantly
so, due to an associated large decrease in the mean condition of non-energy
stressed individuals. The consistently low body condition estimates obtained for
lactating females is in accordance with finding from other studies (Sharp 1982;
Freeland & Choquenot 1990; Gallivan et al. 1995), and reaffirms the high
energetic cost associated with this period of reproduction (Sharp 1982; Gittleman
& Thompson 1988; Clutton-Brock et al. 1989; Randolph & Cameron 2001).
Energy requirements during lactation can be 2.5-5 times that of non-reproductive
females (Gittleman & Thompson 1988). Although there is some evidence of
increased frequency of feeding and food selectivity to offset costs during lactation,
fat deposits prior to breeding or during gestation are thought to be equally, if not
more important for the supplementation of energy needs during lactation
(Gittleman & Thompson 1988). Lactating females are thus expected to drop in
condition as they mobilize their fat reserves to keep up with the demands of
lactation.
46
Old individuals consistently had the lowest body condition estimates (range: 3.3
to 4.5) of all energy classes, with significantly lower conditions than non-energy
stressed individuals in all seasons except winter. Although old individuals had a
mean body condition of 4.5, as compared to 5.6 for non-energy stressed
individuals in winter, the small sample size of old individuals resulted in very
large confidence interval, thereby affecting the sensitivity of the analysis.
Although not significant, Fig. 3.8 illustrates a general decrease in body condition
with age, which is in accordance with findings from other authors (Clutton-Brock
1984; Tyler 1986). The very low body conditions observed in old individuals
highlights the constraints placed upon them by the loss of tooth function (Sinclair
1974; Kojola et al. 1998; Skogland 1988). The tooth surface is critical for the
mechanical breakdown of plant material for further digestion and subsequent
nutrient absorption (Robbins 1983; Fortelius 1985; Skogland 1988; Mackie 2002).
A decrease in the ability to masticate will decrease the amount of soluble cell
content made available for absorption by a decrease in the number of ruptured
cell walls (Owen-Smith 2002). Additionally, it will decrease the amount of surface
area of ingested food exposed to digestion and fermentation in the gut, a process
which is especially important in animals with rapid gut transit times (Mackie 2002).
Despite a low rate, fermentation in the hindgut is likely to cover a large part of the
energy requirements of elephants from the metabolism of volatile fatty acids (Van
Hoven et al. 1981). Both of these factors will thus contribute to a decreased
ability to meet energy and nutrient requirements for the maintenance of condition.
Herbivores occurring at high densities have substantially higher rates of tooth
wear than herbivores occurring at low densities (Skogland 1988; Kojola et al.
1998). This is caused by the inclusion of coarser, lower quality forage due to
competition for limited resources. The elephants of the AMC occur at densities
far above the recommended carrying capacity for their habitat, and it is possible
that the natural loss of tooth function with age could be accelerated in the park,
resulting in further depressed body condition scores.
47
Thus I can surmise that energy stressed elephant life stages are more vulnerable
to resource limitations than non-energy stressed life stages. Specifically, I find
support for the hypotheses that both lactating and old individuals are more
vulnerable to resource limitations than non-energy stressed life stages, whilst
rejecting the hypothesis that weaned individuals are more vulnerable to resource
limitations than non-energy stressed life stages.
3.5.2 Condition and season

Seasonal samples are limited to a single replicate (season) for each of winter,
spring, summer and autumn due to the scope of the study. The following
discussion should therefore be recognized to be related only to these sampled
periods.
Seasonal comparisons revealed that all classes except lactating females

followed the same pattern of a large decrease in condition from winter to spring,
followed by a small increase from spring to summer, and another drop in
condition from summer to autumn. Lactating females exhibited a drop in
condition from both winter to spring and spring to summer, followed by no
change in condition from summer to autumn.
This pattern seems to be correlated with the rainfall recorded in the park
(Appendix 7.5). The higher conditions estimates observed in winter follows a
peak in the bimodal rainfall pattern, with good rains recorded in March (61.2 mm),
April 2007 (31.6 mm) and May 2007 (31 mm). The large drop in condition
observed in spring follows low rainfall in July (15.4 mm), August (16.4 mm) and
September (3.6 mm). The increase in condition observed in summer follows the
second peak in the bimodal rainfall pattern, with good rains recorded in
November (29 mm), December (76.2 mm) and January 2008 (47.6 mm). The
decrease in condition from summer to autumn is again associated with slightly
lower rainfall recorded in February (30.6 mm), March (0 mm) and April 2008
(25.6 mm). Changes in primary productivity in response to rainfall are well
48
recorded (Sinclair 1974; Coe et al. 1976; Phillipson 1975; Owen-Smith 1990;
Mduma et al. 1999). Although no distinct wet and dry season can be
distinguished, Valley Bushveld is known to respond to rainfall with associated
changes in vegetation quality (Hall-Martin et al. 1982). The pattern of body
condition observed thus corresponds to changes in the quality of available
vegetation through the effect of rainfall on primary productivity.
The similarity in responses to seasonal changes across life stages indicate that
both energy stressed and non-energy stressed life stages are sensitive to
changes in resource availability. The non-conformity of lactating females to this
pattern, with a continued decrease in condition, again highlights the extreme
energetic requirements of this stage of reproduction.
My data do not allow a generalization across all seasons/years due to the lack of
replication. These data do, however, suggest that during periods of decreased
primary productivity (due to lower rainfall) elephants can be expected to show a
decline in body condition.
I can thus accept the hypothesis that elephant body condition varies seasonally
with changes in resource availability, and that these seasonal changes in
resource availability will be affected by life stage, with energy stressed life stages
showing a greater response to seasonality than non-energy stressed life stages.
49
CHAPTER 4
SITE-SPECIFIC VARIATION IN BODY CONDITION
¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯
4.1 Introduction
Elephant body condition is not only a reflection of intrinsic factors, but also
extrinsic factors, particularly forage quality and quantity. Understanding the role
of such factors requires a more regional focus to include elephant populations
that experience a range of site-specific forage resources. Furthermore, as is
predicted by Caughley’s (1976b) model, elephants are able to influence forage
resources locally, and this is expected to feedback to influence their body
conditions. This chapter addresses these issues by comparing body condition
across a range of sites, varying in productivity (rainfall), amount of available
resources (reflecting the size of site) and extent of elephant impact on available
resources.
4.1.1 Spatial variation in forage resources

Localized differences in quality and quantity of available forage are well recorded
(Coe et al. 1976; Malpas 1977; Seydack & Bigalke 1992; Seydack et al. 2000).
These landscape-level differences in quality and quantity are generally
associated with variations in rainfall (Sinclair 1974; Coe et al. 1976; Phillipson
1975; Owen-Smith 1990; Mduma et al. 1999; Shackleton 1999; Scholes et al.
2002) and soil nutrients (Coe et al. 1976; Seydack & Bigalke 1992; Grant et al.
1995; Seydack et al. 2000). Forage of higher quality and quantity is generally
associated with areas of higher rainfall and nutrient rich soils.
4.1.2 Impact of herbivores on forage resources

Although landscape differences in forage resources are essentially driven by
large scale factors such as rainfall and soil nutrients, herbivores are able to
impact their habitat and alter patterns of quality and quantity. Overgrazing and
subsequent erosion of soil by wind and water may result in localized depletion of
50
soil nutrients in previously nutrient-rich environments (Grant et al. 1995; Mills &
Fey 2003; Lechmere-Oertle 2005), causing desertification, with associated
declines in productivity (Verstraete & Schwartz 1991).
The occurrence of herbivores at high densities also directly influence the

abundance, and ultimately the quality of available forage through inter-and
intraspecific competition for the shared resource base (Sinclair 1974; Kie et al.
1980; Choquenot 1991; Swihart et al. 1998; Mduma et al. 1999; Van Der Waal &
Grant 2003; Stewart 2005). This effect may be exacerbated by seasonal changes
in plant growth due to changes in rainfall, with density-dependent effects more
apparent during the dry season when nutritious plant growth is limited (Sinclair
1974; Owen-Smith 1990; Van Der Waal 2003). Such an example of severe
depletion of the resource base by high herbivore density can be seen in the
Tsavo elephant die-off of 1970/71, when a prolonged drought caused
exceptionally high aggregations of elephants around waterholes, and subsequent
mortality (Corfield 1973).
The confinement of herbivores within reserves has also created an additional

problem. Although the size of the reserve ultimately determines the number of
herbivores it can support, it also determines the spatial extent for herbivore
movement. This is especially important for larger herbivores that naturally range
over large areas (Owen-Smith 1988). In open systems, elephants tend to roam
over large areas, utilizing different regions at different times of the year (Van
Aarde et al. 2006; O’Connor et al. 2007). When they are confined to small
reserves, this natural movement pattern may be disrupted, forcing them to
remain within the same areas for most of the year (Van Aarde et al. 2006; 2007).
Elephants contained in small reserves tend to have smaller home ranges, with
greater overlap between them (Van Aarde et al. 2007). This causes impacts to
be concentrated over space and time, not allowing vegetation to recover, and
ultimately affecting the resource quality and quantity available (Van Aarde et al.
2006).
51
In addition to the above, the length of exposure to herbivore browsing (i.e. years)
may influence forage resources. Studies in Addo demonstrated that canopy
height and cover, as well as plant species richness showed marked decreases
with increasing length of exposure to elephant utilization (Penzhorn et al. 1974;
Barratt & Hall-Martin 1991; Lombard et al. 2001). Penzhorn et al. (1974) found
that plant biomass was reduced by more than half after 17-18 years of elephant
exposure, with food species being the most affected. According to the predictions
of the exponential model of Lombard et al. (2001), the number of species found
within the Spekboomveld units of the reserve was halved approximately every
seven years. Results showed that over 50% of remnant flora in exposed units
were confined to small populations after 42 years of grazing. Unless elephant
densities were drastically decreased, these trends were predicted to continue.
4.1.3 Effects on body condition

The body condition of an animal is fundamentally governed by the quality and
quantity of the food available (Price & White 1985; Brochu et al. 1988; Pettorelli
et al. 2005). As the quality of food is determined by these various factors (i.e.
rainfall, soil nutrients and herbivore density and impacts), a period of inadequate
supply of any one of these nutrients to meet minimum requirements for
maintenance (and growth in non-mature individuals) will result in loss of body
condition. This has been demonstrated for phosphorus (Wallis de Vries 1996),
nitrogen (Sinclair 1974; Grant et al. 1995) and dietary fibre (Erasmus et al. 1978).
4.1.4 Diet quality and its assessment

Unlike the classical models of optimal diet (Emlen 1966; Schoener 1971), it has
been proposed that large generalist herbivores should not feed to maximize
energy intake per unit time, but rather to optimize the nutrient mix within a given
bulk of food (Westoby 1974). The model proposes that an animal’s dietary choice
is constrained by its need for a certain minimum amount of various nutrients, as
well as the overall limit on the amount of food it can process (obtain or digest)
within a given space of time - thus requiring it to balance the different nutritional
52
benefits against the various costs, rate restrictions and various diet components.
Another such model of an optimal diet under nutrient constraints was proposed
by Pulliam (1975). These models seem far more appropriate for herbivores (as
compared to carnivores), as their food may vary considerably in terms of nutrient
content and digestive constraints over space and time (Crawley 1983).
Furthermore, according to Milton (1979), herbivores should be strongly
influenced by nutritional features, such as the quality or secondary compounds
(antiquality) of potential foods.
The nutritional quality of ingested food is determined by its composition in terms

of the proportion of cell wall to cell contents, by the concentrations of protein,
soluble carbohydrates, minerals and other nutrients in the cell contents, as well
as the degree of lignification of the cell wall (Owen-Smith 2002; O’Connor et al.
2007). Additionally, many plants contain secondary compounds which act to
reduce the nutrients available to herbivores (Robbins 1983; Robbins, Mole,
Hagerman & Hanley 1987; Owen-Smith 2002) or have toxic effects (Milton 1979).
Protein is regarded as one of the most important and often limiting components
of wildlife diets. The continuous availability of dietary nitrogen, an estimate of
crude protein (% N x 6.25), is essential for the growth and maintenance of all
animal tissues (Robbins 1983). Faecal nitrogen as an indicator of diet quality has
been used in a number of herbivore studies (Sinclair 1974; Mould & Robbins
1981; Hall-Martin et al. 1982; Wofford et al. 1985; Leslie & Starkey 1987; Grant
et al. 1995; Wrench et al. 1997; Van der Waal et al. 2003; Osborn 2004), and has
been found to be correlated to dietary protein (Mould & Robbins 1981; Wofford et
al. 1985; Howery & Pfister 1990; Grant et al. 1995; Wrench et al. 1997).
It is not adequate, however, to explain dietary quality only in terms of these

components. The dietary fibre component is equally important in determining the
quality of forage. “Dietary fibre” is often used to refer to the plant cell wall, which
consists mainly of cellulose and hemicellulose (Robbins 1983). These
53
components are not susceptible to enzymatic digestion in vertebrates, but can be
broken down by gastrointestinal flora found in the digestive tracts of most
herbivorous animals (Robbins 1983). Herbivores have a tendency to selectively
feed on less fibrous plants and plant parts (Bell 1971; Owen-Smith & Cooper
1987; Cooper et al. 1988). Possible reasons for this tendency include:
1) Cell wall components prevent the extraction of cell nutrients (Keys et al.
1969; Bell 1971; Janis 1976), therefore the more fibrous the food, the less
available the nutrients within cells.
2) Fibre and lignin content has been negatively correlated with digestibility
in ruminants and non-ruminants (Schneider et al. 1951; 1952; Owen-Smith
2002; Hatt et al. 2005).
A high fibre content is therefore associated with low quality forage (Erasmus et al.
1978; Demment & Van Soest 1985; Owen-Smith 1988). Additionally, a high fibre
content is generally associated with a low protein content, as protein
concentration is largely determined by the cell contents to cell wall ratio
(Erasmus et al. 1978; Owen-Smith 2002).
In addition to these requirements, herbivores also need a variety of minerals to

maintain cellular activity and body functions (Robbins 1983). Furness (1988)
drew attention to the role of mineral nutrition on diet selection by herbivores,
which lead to the suggestion that minerals should be included in optimal foraging
models, especially relating to seasonal requirements (Bazely 1989). Deficiencies
or imbalances of minerals play an important role in determining animal condition,
fertility, productivity and mortality, and it is thought that marginal deficiencies may
be easily overlooked in favour of more obvious factors causing population
fluctuations, such as severe weather, food shortages, parasites or infectious
agents (Underwood 1977). Of all essential minerals, sodium and phosphorus are
increasingly being shown to be the two minerals governing the body condition
and diet selection of large herbivores when soil fertility is primarily low
(McNaughton 1988; 1990; Freeland & Choquenot 1990; Ben-Shahar & Coe
1992). Both are essential for normal growth and reproduction (Weir 1972;
54
Groenewald & Boyazoglu 1980; Grassman & Hellgren 1993; Ulrey et al. 1997;
Wrench et al. 1997; Dörgeloh et al. 1998; Seydack et al. 2000). According to the
Department of Environmental Affairs and Tourism’s Eastern Cape state of the
environment report (DEAT 2004), a large portion of the Eastern Cape has a
naturally high salinity in both its ground and surface water. As elephants are
known to drink large volumes of water on a daily basis (Weir 1972), it is thus
unlikely that sodium would prove to be a limiting nutrient in this area. A previous
study by Koen et al. (1988) on the macronutrients available to elephants in the
AENP has noted, however, that the phosphorus levels in both the plants species
and faeces analysed was low, with a very high calcium: phosphorus ratio, which
would hamper the absorption of the little phosphorus available, suggesting that
phosphorus levels may affect the growth and reproduction of elephants in this
region.
4.1.5 Elephant nutrient requirements

According to McCullagh (1969b), a young growing elephant with a body weight of
1000 kg would require c. 0.3 kg of digestible protein on a daily basis. Malpas
(1977) estimated that this would require an average of 6% crude protein in the
diet, and showed that elephant body condition estimates from Uganda support
this figure. More recently, Ulrey et al. (1997) proposed minimum nutrient
requirements for African elephants based largely on extrapolations from the
nutrient requirements of horses. They estimated that adult elephants would
require a minimum of 8% crude protein for maintenance, 10-12% crude protein
for pregnancy, and 12-14% during lactation, whilst 4-12 year olds were estimated
to require 12% crude protein.
Wrench et al. (1997) estimated maintenance phosphorus requirements at 0.2%

for most ungulates. This was supported by Ulrey et al. (1997), who also
estimated phosphorus requirements at 0.2% for maintenance of elephants, with
this value increasing to 0.3% for pregnancy and lactation, and 0.3% for 4-12 year
olds.
55
As elephants are large-bodied hindgut fermenters, they are able to persist on a
diet with a very high fibre content. Meissner et al. (1990) found an average of
62% NDF in the diet of elephant in the Kruger National Park.
4.2 Hypotheses and aims

By investigating the body condition of elephant populations occurring at various
sites differing in rainfall, size, occupational history and density, I am able to test
the hypothesis that elephant body condition will vary as a function of primary
productivity. Specifically, I predict that there will be a decline in overall body
condition at sites with low rainfall, high elephant density, long elephant
occupational history and small size.
By investigating the body condition of elephants across various sites in relation to

the dietary quality available to them, I am able to test the hypothesis that
elephant body condition will vary as a function of resource quality. Specifically, I
predict that there will be a decrease in overall condition at sites with low levels of
dietary protein and phosphorus and high levels of NDF.
This will enable me to determine whether elephant body condition is sensitive to

site-specific differences in resources and allow me to identify populations that
may be resource limited.
4.2 Materials and methods

4.3.1 Choice of sampling method
Four different methods of obtaining samples for the determination of diet quality
of herbivores exist:
1) Observing the animals directly and then attempting to sample those
plants and plant parts taken for dietary quality analysis (e.g. Meissner et al.
1990; Wallis de Vries 1996). This method is very labour intensive and
requires tame animals.
56
2) Inserting oesophageal fistulas in the animals (Engels et al. 1971). This
usually requires tame animals and is only practical for a limited number of
herbivores.
3) Obtaining a rumen or stomach sample from the animals (e.g.
McCullagh 1969b; Malpas 1977). This requires tame animals to insert a
rumen fistula or killing the animals to obtain the samples.
4) Collecting fresh faecal samples and using that as representative of the
forage taken (e.g. Dörgeloh et al. 1998; Van Der Waal & Grant 2003). This
method does not require tame or surgically-fitted animals and does not
involve killing the animals.
Of all these methods, the collection of faecal samples is the most practical for the
study of wild herbivores (Erasmus et al. 1978), and was therefore used here.
4.3.2 Measures of dietary quality

Although there has been some debate on the value of faecal indices as
indicators of dietary quality (Hobbs 1987), it has been found that faecal nitrogen
is correlated with dietary nitrogen, and could thus be used to assess seasonal
changes in dietary quality of a single population as well as within-season
comparisons of different populations that occupy like habitats (Leslie & Starkey
1987). Faecal phosphorus levels have also been found to correlate to dietary
phosphorus levels (Moir 1966; Howery & Pfister 1990; Wrench et al. 1997).
Additionally, these correlations are improved by defining the season and locality,
as well as by considering faecal nitrogen and phosphorus levels together, as
their excretion is linked (Moir 1966).
NDF represents the total fibre component of the plant, and is thought to more
closely correspond to the indigestible cell wall fraction than the more widely used
crude fibre estimates (Ulrey et al. 1997), and has been shown to be a useful
measure of diet quality (Hall-Martin et al. 1982).
57
4.3.3 Sampling procedure
Each of the 6 study sites (see Chapter 2, same abbreviations) was visited once
per season between July ’07 and July ’08 (winter = July/Aug 2007; spring =
Oct/Nov 2007; summer = Jan/Feb 2008; autumn = April/May 2008), except for
Shamwari, which was not visited during spring. For each site and season, visual
body condition scores (as described in Chapter 2) were recorded (see Appendix
1 for sample sizes), and 5 fresh (<2 days old) faecal samples collected and
placed in labelled brown paper bags. Additionally, fresh plant clippings were
obtained from the AMC and NCA seasonally, and from all other sites for summer
and winter. Plant clippings were taken from at least three plants per species per
season at different locations for each reserve to account for interplant variation.
The plants collected formed part of the known elephant diet for each site,
obtained from previous dietary studies for Asante Sana (Minnie 2006), Addo
(Landman et al. 2007), Nyathi (Davis 2005), Shamwari (Roux 2006) and Kariega
(Wolmarans 2006), and from direct observations by reserve managers for
Blaauwbosch. All the principal dietary items (PDI’s; those species contributing
3% or more to the diet) identified for AMC and NCA were collected, but only
overlapping dietary items were collected from the other reserves. These were
plant species that formed part of the known elephant diet for at least three of the
sampled sites, and included Azima tetracantha, Schotia afra, Cynodon dactylon
and Acacia karroo.
All samples collected were placed in labelled brown paper bags and oven dried
at 60°C for a minimum of 3 days. All plants of each species per season were
added together to create one sample per species per season. These samples
were then analysed at the Grootfontein Agricultural Development Institute,
Middleburg, Eastern Cape. Samples were ground to 2 mm mesh size,
homogenised and subsets of each sample were then analyses for phosphorus
(P), nitrogen (N) and neutral detergent fibre (NDF). Nitrogen was analysed using
the Kjeldahl method as set out by the Association of Official Agricultural
Chemists (AOAC 1995), whilst NDF was determined by the Ankom Technology
58
method filter bag technique (www.ankom.com) and phosphorus by the dry ashing
method (ultraviolet-visible spectrophotometry, AOAC 1995). Nitrogen values
were subsequently converted to crude protein (%N x 6.25) by standard practice
(Robbins 1983). All values are expressed as percentage dry weight. In order to
test the precision of the sample analysis, 5 blind replica samples were submitted
without the knowledge of those doing the analysis.
In addition to the body condition and dietary quality data collected, rainfall (both
long term and for the study period and the preceeding six month time), reserve
size, history of elephant occupation and elephant density data were collected at
each site. Where rainfall data were not recorded on site, the nearest
town/weather station rainfall data were used. Study period rainfall was recorded
as rainfall over the six month prior the commencement of the study to six months
before the end of the study to allow a more accurate representation of vegetative
responses in productivity. Unless stated otherwise, where rainfall during the
study period is mentioned, it refers to this recorded rainfall.
4.3.4 Data analysis

4.3.4.1 Body Condition
As body conditions in the initial AMC analysis (Chapter 3) were found to differ
significantly between life stages, a heterogeneity chi-squared test (Zar 1999) was
performed for each season to determine whether energy stressed life stages (i.e.
old age, lactating and weaning) could be pooled for subsequent body condition
comparisons with other sites. As body conditions in the initial AMC analysis also
were found to differ significantly between seasons, these data, and those from
other sites, were kept separate in subsequent analyses.
A four-way log linear analysis (Statistica 8.0; StatSoft Inc. 2007) was used to
determine whether there was a significant difference in body condition between
the various sites and seasons. Factors were site, season, energy class (non-
energy stressed and energy stressed) and body condition category. Data were
59
then explored graphically using Box-and-Whiskers plots (Statistica 8.0; Statsoft
Inc. 2007), with whiskers representing the 90% confidence interval (CI).
A proportional odds ordinal logistic regression model was used to investigate

which site specific factors (i.e. rainfall, density, reserve size and occupational
history) would best explain the body conditions observed. Sites were weighted by
the relative frequency of occurrence of each body condition class observed.
Backward stepwise selection was used to select the most parsimonious model.
This process involved first fitting the model with all variables, removing non-
significant variables, and re-fitting the model using only the significant variables
(elephant density and rainfall over the course of the study period). Model
goodness-of-fit was assessed using Nagelkerke’s R2 (Nagelkerke 1991). The
proportional odds logistic regression model was fitted using the package “Design”
(Harrell 2008) on the statistical platform R (R.D.C.T. 2008).
4.3.4.2 Dietary quality

A paired t-test (Statistica 8.0; StatSoft Inc. 2007) of the blind replicate samples
was used to determine the precision of the dietary quality analyses obtained from
the Grootfontein Agricultural Institute.
The three components (protein, phosphorus and NDF) of the faecal dietary
quality data were then analysed using factorial ANOVAs (Statistica 8.0; StatSoft
Inc. 2007) to investigate whether any differences existed between the seasons
and sites. Significant results were further analysed using Tukey’s post hoc tests
in order to explore the significant interactions. Subsequently, a proportional odds
ordinal logistic regression model was used to investigate which of the dietary
quality measures best explained the ordering and observed body conditions. The
best model was again selected using the backward stepwise method (as
explained in 4.3.4.1) and goodness of fit was assessed using Nagelkerke’s R2
(Nagelkerke 1991). The proportional odds logistic regression model was fitted
60
using the package “Design” (Harrell 2008) on the statistical platform R (R.D.C.T.
2008).
The various diet quality components of overlapping dietary plant species were
analysed using paired t-tests (Statistica 8.0; StatSoft Inc. 2007) to investigate any
differences between seasons. The dietary quality values obtained were also
explored graphically to investigate any differences between sites. Two-way
ANOVAs (Statistica 8.0; StatSoft Inc. 2007) were used to determine whether any
differences in quality between the PDI’s of the AMC and NCA could be detected.
Where data were not normally distributed, log10-transformations were done to

allow conformation with the requirements of parametric statistics. Where data
were log transformed for statistical analyses, the data were back transformed to
report results in the original units. Except where stated otherwise, tests were
assessed at the 95% probability level (Zar 1999).
4.4 Results
4.4.1 Body Condition
Heterogeneity chi-squared analyses revealed that there was no difference in the
body conditions of the three energy stressed classes within seasons (winter: χ2 =
7.02, df = 6, p = 0.32; spring: χ2 = 7.66, df = 6, p = 0.26; summer: χ2 = 9.40, df =
6, p = 0.15; autumn: χ2 = 12.01, df = 6, p = 0.06), and that these classes could
thus be pooled within seasons for subsequent analyses.
From the log-linear analysis of body condition, it was found that no three-way
interactions were significant, and that the data were adequately explained by
two-way interactions (χ2 = 359.77, df = 50, p < 0.001). The model that best
explained the results included the interactions (1) site x season, (2) site x energy
class, (3) site x body condition category, (4) season x body condition category
and (5) energy class x body condition category. However, as the factor of
61
principal interest was the body condition category, interactions 1 and 2 were not
further explored.
Thus the results indicate that, controlling for season and energy class, body
condition was not the same for all sites (χ2 = 184.67, df = 15, p < 0.001).
Controlling for site and energy class, body condition also was found to differ
significantly between seasons (χ2 = 50.55, df = 9, p < 0.001). A significant
interaction also was found between body condition and energy classes,
controlling for site and season (χ2 = 16.05, df = 3, p = 0.001). See Appendix 3 for
full results.
Graphical exploration of these results (Fig. 4.1) indicated that in winter energy
stressed elephants in the AMC were in poorer condition (BC = 4.8) than those
from the other reserves, with this difference being significant for NCA (BC = 5.8),
Asante Sana (BC = 5.8) and Shamwari (BC = 5.5). Comparisons for non-energy
stressed individuals for the same season revealed no significant differences
between sites (Fig. 4.2).
In spring, energy stressed individuals were in significantly poorer condition in the

AMC (BC = 4.3) than in any of the other reserves. Individuals in Asante Sana
(BC = 6.0) also had significantly better condition than those from the other
reserves (Fig. 4.3). Non-energy stressed individuals from the AMC were also in
poorer condition (BC = 4.7) than those from other reserves, although this
difference was only significant for Blaauwbosch (BC = 6.0) and Asante Sana (BC
= 5.8) (Fig. 4.4).
62
6.6
6.4
6.2
6.0
5.8
Body Condition
5.6
5.4
5.2
5.0
4.8
4.6
4.4
AMC Kariega Blwbosch
NCA ASana Shamwari
Fig. 4.1 Mean body condition, ± 0.9 confidence interval, for energy stressed (i.e.
lactating, weaned and old age) elephants in winter. AMC = Addo Main Camp,
NCA = Nyathi Concession Area, Asana = Asanta Sana, Blwbosch =
Blaauwbosch.
6.4
6.2
6.0
5.8
Body Condition
5.6
5.4
5.2
5.0
4.8
4.6
NCA ASana Shamwari
Fig. 4.2 Mean body condition, ± 0.9 confidence interval, for non-energy stressed
(i.e. sub-adult and non-lactating) elephants in winter. Sites as per Fig 4.1.
63
6.2
6.0
5.8
5.6
5.4
Body Condition
5.2
5.0
4.8
4.6
4.4
4.2
4.0
AMC NCA Kariega ASana Blwbosch
Fig. 4.3 Mean body condition, ± 0.9 confidence interval, for energy stressed
elephants in spring. Sites as per Fig 4.1.
6.4
6.2
6.0
5.8
5.6
Body Condition
5.4
5.2
5.0
4.8
4.6
4.4
4.2
AMC NCA Kariega ASana Blwbosch
elephants in spring. Sites as per Fig 4.1.
A similar pattern was observed for summer, with energy stressed AMC elephants
exhibiting significantly lower body conditions (BC = 4.2) than those from other
64
reserves. The highest body condition scores for this class were obtained for
Blaauwbosch (BC = 5.7), which had significantly better condition than all
reserves, with the exception of Kariega (BC = 5.4) (Fig. 4.5). Non-energy
stressed individuals in the AMC again had lower scores (BC = 4.8) than those
animals from other reserves, with this difference being significant for Kariega (BC
= 5.5), Asante Sana (BC = 5.8) and Shamwari (BC = 5.4) (Fig. 4.6).
In autumn, energy stressed individuals in the AMC again exhibit significantly

lower body conditions (BC = 4.1) than those from all other reserves (Fig. 4.7).
Energy stressed elephants from the NCA were also in poorer condition (BC =
4.9), with this difference being significant for Blaauwbosch (BC = 5.7) and
Shamwari (BC = 5.4). Non-energy stressed animals from the AMC follow the
same trend as the energy stressed individuals in autumn, with significantly lower
body condition estimates (BC = 4.4) than other reserves (Fig. 4.8). No
differences in condition were observed for non-energy stressed individuals
between the other reserves.
6.2
6.0
5.8
5.6
5.4
Body Condition
5.2
5.0
4.8
4.6
4.4
4.2
4.0
3.8
NCA ASana Shamwari
Fig. 4.5 Mean body condition, ± 0.9 confidence interval, for energy stressed
elephants in summer. Sites as per Fig 4.1.
65
6.4
6.2
6.0
5.8
Body Condition
5.6
5.4
5.2
5.0
4.8
4.6
4.4
NCA ASana Shamwari
elephants in summer. Sites as per Fig 4.1.
6.2
6.0
5.8
5.6
5.4
Body Condition
5.2
5.0
4.8
4.6
4.4
4.2
4.0
3.8
3.6
NCA ASana Shamwari
Fig. 4.7 Mean body condition, ± 0.9 confidence interval, for energy-stressed
elephants in autumn. Sites as per Fig 4.1.
66
6.4
6.2
6.0
5.8
Body Condition 5.6
5.4
5.2
5.0
4.8
4.6
4.4
4.2
4.0
NCA ASana Shamwari
elephants in autumn. Sites as per Fig 4.1.
Energy stressed individuals in the AMC consistently exhibit body condition

scores in the range of 4.0 - 4.8 (“fair” according to the body condition scale of
Gough (in prep.)), almost consistently significantly lower than similar individuals
from other reserves. Although not significant for all seasons, non-energy stressed
individuals showed a similar pattern, with generally lower condition scores than
non-energy stressed individuals from other reserves.
Although the AMC was the source of most variation between sites, some
individual differences in condition between other sites were also detected, as
highlighted above (Figs 4.3, 4.4 and 4.5).
In the AMC, body condition was significantly higher in winter for both energy
stressed and non-energy stressed classes than in any other season (Fig. 4.9).
Both energy classes followed a general downward trend in body condition from
winter to autumn, with both experiencing a significant drop in condition from
summer to autumn (from 5.6 to 4.7 for non-energy stressed, and from 4.8 to 4.3
for energy stressed individuals). Within seasons, energy stressed individuals
67
were always found to be in significantly poorer condition than non-energy
stressed individuals, except in spring when this difference was not significant.
In the NCA, body condition was significantly higher in winter (5.8) for energy
stressed individuals than any other season (Fig. 4.10). Non-energy stressed
individuals also had higher condition estimates in winter (5.7) than other seasons,
but this was not significant. Both energy classes experienced a general
downward trend in condition from winter to autumn, with non-energy stressed
individuals showing a slight increase from spring to summer. No significant
difference in condition was found between the energy classes within seasons,
although non-energy stressed individuals exhibited better condition than energy
stressed individuals in all seasons but winter.
6.0
5.8
5.6
5.4
5.2
Body Condition
5.0
4.8
4.6
4.4
4.2
4.0
3.8
WinterES SpringES SummerES AutumnES
WinterNES SpringNES SummerNES AutumnNES
Fig. 4.9 Seasonal mean body condition, ± 0.9 confidence interval, for energy
stressed and non-energy stressed elephants in the Addo Main Camp. Capital
letters of X-Axis labels represent energy class (i.e. ES = energy stressed, NES =
non-energy stressed).
68
6.2
6.0
5.8
5.6
Body Condition
5.4
5.2
5.0
4.8
4.6
4.4
stressed and non-energy stressed elephants in the Nyathi Concession Area.
Although some minor changes in condition were exhibited, elephants in Kariega

experienced no significant changes in condition for either energy classes
between seasons (Fig. 4.11). There was also no significant difference in
condition detected between the two energy classes, although energy stressed
individuals tended to be in worse condition than non-energy stressed individuals.
This held for all seasons except spring.
In Asante Sana, no significant change in elephant condition was observed for

non-energy stressed individuals (Fig. 4.12). Energy stressed individuals showed
a slight increase in condition from winter to spring, followed by a significant drop
in condition from spring (BC = 6.0) to summer (BC = 5.0), and again a slight
increase from summer to autumn. In winter and spring, energy stressed
elephants were in slightly better condition than non-energy stressed ones, with
this pattern being reversed in summer and autumn, although none of these
differences were significant. Although some points exhibited very small
confidence intervals due to zero variation among observations, the generally
69
large confidence intervals were a result of the small population size (n = 9), and
affected the sensitivity of the results.
6.2
6.0
5.8
Body Condition
5.6
5.4
5.2
5.0
4.8
stressed and non-energy stressed elephants in Kariega.
6.6
6.4
6.2
6.0
5.8
Body Condition
5.6
5.4
5.2
5.0
4.8
4.6
4.4
stressed and non-energy stressed elephants in Asante Sana.
70
No significant differences in seasonal condition could be detected in
Blaauwbosch (Fig. 4.13) for energy stressed individuals, whilst non-energy
stressed individuals were in significantly better condition in spring (BC = 6.0) than
in winter (BC = 5.4) or summer (BC = 5.3). Within seasons there was no
significant difference in condition estimates for the two energy classes, except for
spring, in which non-energy stressed individuals were in significantly better
condition.
In Shamwari (Fig. 4.14), energy stressed individuals experienced a significant

decrease in condition from winter (BC = 5.5) to summer (BC = 4.7), followed by a
significant increase from summer to autumn (BC = 5.4). Non-energy stressed
individuals showed no difference across seasons. No differences in condition
between the two energy classes could be detected within any season.
6.4
6.2
6.0
5.8
Body Condition
5.6
5.4
5.2
5.0
4.8
4.6
WinterNES SpringNES SummerNES AutumnNE
Fig. 4.13 Season mean body condition, ± 0.9 confidence interval, for energy
stressed and non-energy stressed elephants in Blaauwbosch.
71
6.0
5.8
5.6
5.4
Body Condition
5.2
5.0
4.8
4.6
4.4
WinterES SummerES AutumnES
WinterNES SummerNES AutumnNES
Fig. 4.14 Seasonal mean body condition for energy stressed and non-energy
stressed elephants in Shamwari.
Table 4.1 Potential environmental and site-specific drivers of elephant body

condition from six sites in the Eastern Cape.
Period of
Area Density Long term Study period
SITE occupation
(km2) (km2) average rain rainfall
(yrs)
Addo Main Camp 103 4.029 396 364 54
Nyathi Concession Area 114 0.781 396 364 5
Blaauwbosch 45 0.222 300 332 3
Kariega 190 0.100 640 684 4
Shamwari 180 0.261 550 427 15
Asante Sana 108 0.083 360 645 4
The full proportional odds ordinal logistic regression model revealed that of the
environmental drivers (Table 4.1), only study period rainfall and elephant density
had significant p values for inclusion in the model (see Appendix 4 for full results
& Appendix 7 for rainfall distribution during the study period), with an R2 value of
0.36. The reduced model with only rainfall over the study period and density had
an R2 value of 0.354, indicating that reducing the model down to two parameters
did not reduce the goodness-of-fit and that the other parameters are therefore
fairly redundant.
72
When the proportion of the population is plotted against the population density for
each body condition score (Fig. 4.15), larger proportions of the two worst body
conditions scores (BC 3 & 4) are associated with high population densities,
whereas the proportions of the two best body condition scores (BC 5 & 6) are
shown to decrease with increasing population densities. These relationships are
significant for body condition 3 (p < 0.001), 4 (p < 0.001) and 6 (p = 0.014).This
confirms the results obtained in the proportional odds model.
0.7
0.6
r2 = 0.979
Proportion of the Population
0.5
0.4
r2 = 0.346
0.3
0.2
0.1 r2 = 0.991
r2 = 0.816 BC 3
BC 4
0.0
BC 5
-0.5 0.0 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5
BC 6
Density
Fig. 4.15 The relationship between density of elephants and body condition,
expressed as a proportion of the population for each body condition score.
Although the relationships are not significant, the proportion of individuals with
the best body condition score (BC 5 & 6) is higher in reserves with high rainfall
during the study period, with those in the worst body condition (BC 3 & 4) being
proportionally more present in reserves with low rainfall during the study period
(Fig. 4.16).
73
0.7
0.6
r2 = 0.304
Proportion of population 0.5
r2 = 0.006
0.4
0.3
0.2
r2 = 0.213
0.1
r2 = 0.131
BC 3
BC 4
0.0
BC 5
300 350 400 450 500 550 600 650 700
BC 6
Rainfall during study
Fig. 4.16 The relationship between rainfall during the study period and elephant
body condition, expressed as a proportion of the population for each body
condition score.
4.4.2 Dietary quality

4.4.2.1 Precision of sample analysis
Paired t-tests revealed no significant differences between results obtained for
samples and their blind replicates for protein (p = 0.20; df = 4), phosphorus (p =
0.54; df = 4) or NDF (p = 0.35; df = 4). Results obtained from analyses could thus
be considered to be a precise reflection of the actual values.
4.4.2.2 Faecal dietary quality

From the ANOVA, faecal protein was found to differ significantly between sites (F
= 2.97, df = 4, p = 0.023) and seasons (F = 34.43, df = 2, p < 0.001). The
interaction between season and site was also found to be significant (F = 2.84, df
= 14, p = 0.001). See Appendix 5 for full results.
For within season, between site comparisons of faecal protein (Table 4.2),
although slight differences were detected, Tukey’s post hoc revealed no
74
significant differences between sites for winter, with Asante Sana (4.91%),
Blaauwbosch (5.55%) and the AMC (5.50%) having the lowest values, and
Shamwari the highest (7.84%). Spring revealed a significant difference in faecal
protein values between the AMC (4.29%) and the NCA (7.56%). No difference in
faecal protein values were observed during summer. Autumn revealed
significantly higher faecal protein values for Shamwari (9.88%) than either the
AMC (6.73%) or NCA (5.94%).
Table 4.2 Mean percentage faecal protein for each season for comparisons
between sites. Letters denote significant differences among means as
established by Tukey’s post hoc test. n/s = not sampled.
SITE WINTER % SPRING % SUMMER % AUTUMN %
Addo Main Camp 5.55 a 4.29 a c 8.55 a 6.73 a c
Nyathi Concession
Area 6.53 a 7.56 b 7.23 a 5.94 a c
Kariega 6.30 a 6.56 a 9.55 a 7.83 a
Asante Sana 4.91 a 6.94 a 9.05 a 7.65 a
Blaauwbosch 5.50 a 6.60 a 9.90 a 7.41 a
Shamwari 7.84 a n/s 9.08 a 9.88 b
The ANVOA of faecal phosphorus indicated that faecal phosphorus also differed
significantly between sites (F = 6.40, df = 4, p < 0.001) and seasons (F = 26.37,
df = 2, p < 0.001), and that the interaction between season and site was
significant (F = 3.12, df = 14, p < 0.001). See Appendix 5 for full results.
Tukey’s post hoc for within-season comparisons between sites for faecal
phosphorus (Table 4.3) revealed significantly higher values in Shamwari (0.27%)
than the AMC (0.13%) or Kariega (0.15%) for winter, whilst the three other
reserves all had phosphorus levels only slightly higher than the AMC and Kariega.
In spring the AMC again had the lowest percentage phosphorus (0.12%), and
along with Asante Sana (0.14%), had significantly lower phosphorus contents
than that of the NCA (0.26%), with Kariega and Blaauwbosch having
intermediate values. No difference in faecal phosphorus was detected during
summer, but this changed in autumn, when Shamwari (0.32%) had significantly
higher phosphorus levels than either the AMC (0.16%) or NCA (0.17%).
75
Table 4.3 Mean percentage faecal phosphorus for each season for comparisons
between sites. Letters denote significant differences among means as
established by Tukey’s post hoc test. n/s = not sampled.
SITE WINTER % SPRING % SUMMER % AUTUMN %
Addo Main Camp 0.13 a c 0.12 a c 0.25 a 0.16 a c
Nyathi Concession
Area 0.18 a 0.26 b 0.24 a 0.17 a c
Kariega 0.15 a c 0.18 a 0.26 a 0.24 a
Asante Sana 0.17 a 0.14 a c 0.24 a 0.25 a
Blaauwbosch 0.17 a 0.16 a 0.23 a 0.21 a
Shamwari 0.27 b n/s 0.32 a 0.32 b
For within-site, between season comparisons of faecal protein (Table 4.4),

Tukey’s post hoc test revealed that, in the AMC, faecal protein was significantly
higher in summer (8.55%) than in spring (4.29%), with intermediate values in
autumn and winter. No significant differences in faecal protein were observed in
the NCA. Kariega had significantly higher faecal protein values in summer
(9.55%) than in winter (6.30%). This was also true of Asante Sana (summer =
9.05%, winter = 4.91%). Faecal protein in Blaauwbosch showed a significant
increase from spring (6.60%) to summer (9.90%), with the difference between
winter (5.50%) and summer also being significant. Faecal protein values in
Shamwari revealed no significant changes between seasons.
Table 4.4 Mean percentage faecal protein for each site for comparisons between
seasons. Letters denote significant differences among means as established by
Tukey’s post hoc test.
AMC NCA KARIEGA A SANA BLWBOSCH SHAMWARI
SEASON % % % % % %
Winter 5.55 a 6.53 a 6.30 a c 4.91 a c 5.50 a c 7.84 a
Spring 4.29 a c 7.56 a 6.56 a 6.94 a 6.60 a c n/s
Summer 8.55 b 7.23 a 9.55 b 9.05 b 9.90 b 9.08 a
Autumn 6.73 a 5.94 a 7.83 a 7.65 a 7.41 a 9.88 a
The highest protein values were found in summer for all reserves, except the
NCA and Shamwari, for which spring and autumn had the highest values,
respectively. The lowest protein values were obtained in winter for all reserves
except the AMC and the NCA, for which spring and autumn samples contained
the least amount of protein.
76
Tukey’s post hoc test for faecal phosphorus (Table 4.5) indicated that
phosphorus values in the AMC were significantly higher in summer (0.25%) than
in winter (0.13%) or spring (0.12%). No significant differences in phosphorus
values were observed in the NCA. Kariega had significantly higher protein and
phosphorus values in summer (0.25%) than in winter (0.15%). In Asante Sana,
faecal phosphorus showed a significant increase from spring (0.14%) to summer
(0.24%). Autumn values (0.25%) remained high, and was also significantly higher
than spring. Blaauwbosch showed no significant differences in faecal phosphorus
between seasons. Kariega, Asante Sana and Blaauwbosch all followed a similar
pattern of increasing protein values from winter to spring, and again from spring
to summer, with a subsequent drop in protein from summer to autumn. Faecal
phosphorus levels in Shamwari showed no significant changes across seasons.
Table 4.5 Mean percentage faecal phosphorus for each site for comparisons
between seasons. Letters denote significant differences among means as
established by Tukey’s post hoc test.
SEASON % % % % % %
Winter 0.13 a c 0.18 a 0.15 a c 0.17 a 0.17 a 0.27 a
Spring 0.12 a c 0.26 a 0.18 a 0.14 a c 0.16 a n/s
Summer 0.25 b 0.24 a 0.26 b 0.24 b 0.23 a 0.32 a
Autumn 0.16 a 0.17 a 0.24 a 0.25 b 0.21 a 0.32 a
Summer phosphorus levels were higher than those in winter and spring, except
for the NCA, where the spring average was higher than summer. Autumn values
also were found to be higher than winter and spring, except for the NCA, for
which the autumn average was lower than both winter and spring. Phosphorus
levels in Shamwari were found to be consistently higher than values obtained
from other reserves.
The initial ANOVA of NDF revealed a significant difference between sites, but no
difference between seasons. There was also not a significant interaction between
site and season. Tukey’s post hoc test attributed the significant site result to a
difference in fibre values between the AMC (79.48%) and Asante Sana (66.36%)
in spring. After inspection of the data, it was found that this lower value in Asante
77
Sana was due to an anomalously low value obtained for one of the replicates.
The analysis was subsequently redone without this outlier. The final ANOVA of
NDF revealed a significant difference between sites (F = 3.82, df = 4, p = 0.006),
with seasons also being significantly different (F = 7.15, df = 2, p = 0.001). The
interaction between site and season also was found to be significant (F = 2.07, df
= 14, p = 0.02). See Appendix 5 for full results.
Tukey’s post hoc test (Table 4.6) attributed the seasonal differences to lower
NDF values obtained in the Kariega summer (68.1%) and the AMC summer
(70.8%) samples as compared to their spring values (Kariega = 76.5%, AMC =
79.5%). Tukey’s test did not, however, reveal any significant difference between
sites within seasons. Any other differences were as a result of the interactions
between various seasons of the different sites (e.g. AMC spring significantly
differed from Kariega summer, etc.).
Table 4.6 Mean percentage faecal NDF for each site for comparisons between
seasons. Letters denote significant differences among means as established by
Tukey’s post hoc test.
SEASON % % % % % %
Winter 77.14 a 76.89 a 68.13 a 70.56 a 74.05 a 70.95 a
Spring 79.49 a c 72.02 a 72.07 a c 73.47 a 76.57 a n/s a
Summer 70.82 b 74.58 a 74.28 b 70.77 a 69.33 a 71.05 a
Autumn 76.70 a 75.45 a 76.54 a 66.36 a 71.46 a 70.05 a
The results from the ordinal logistic regression model revealed that the simplest
model that explained the observed body condition distribution was protein and
NDF. The full model had an R2 value of 0.275, whilst the reduced best-fit model
had an R2 value of 0.274, indicating that the parameter removed was redundant
(see Appendix 4 for full results). These R2 values are, however, fairly low and
suggests that the model was not very good fit for the data.
When the proportion of the population within each body condition score are
plotted against the faecal protein values obtained (Fig. 4.17), populations with
larger proportions of individuals in the two best body condition (BC 5 and 6)
78
scores tend to be correlated with higher protein values recorded on those
reserves, and populations with larger proportions of individuals in the two worst
body condition scores (BC 3 & 4) tend to be correlated with lower protein values
recorded on those reserves. However, none of these relationships were found to
be significant.
0.7
0.6
0.5
Proportion of population
r2 = 0.370
0.4 r2 = 0.154
0.3
0.2
r2 = 0.302
0.1
r2 = 0.374 BC 3
BC 4
0.0
BC 5
6.0 6.2 6.4 6.6 6.8 7.0 7.2 7.4 7.6 7.8 8.0 8.2 8.4 8.6 8.8 9.0 9.2
BC 6
Faecal Protein
Fig. 4.17 The relationship between elephant faecal protein and body condition,
expressed as a proportion of the population for each body condition score.
When I plot the proportion of the population within each body condition score
against the faecal NDF values obtained (Fig. 4.18), one can see the reverse
pattern as observed with faecal protein, with the two worst body conditions
showing a proportional increase with an increase in NDF, and the best body
condition showing a proportional decrease with an increase in NDF. Although
these relationships are stronger than those found for protein, the only significant
relationship is that of the proportion of BC 6 and faecal NDF (p = 0.049).
79
0.7
0.6
0.5
Proportion of population
r2 = 0.022
0.4
r2 = 0.660
0.3
0.2
r2 = 0.568
0.1
r2 = 0.530
BC 3
BC 4
0.0
BC 5
70 71 72 73 74 75 76 77
BC 6
Faecal NDF
Fig. 4.18 The relationship between faecal NDF and body condition, expressed as
a proportion of the population for each body condition score.
4.4.2.3 Plant dietary quality

Although paired t-test analyses of protein, phosphorus and NDF found no
significant differences for any of the species in seasonal (winter-summer)
comparisons within sites, some differences were still detectable between
seasons. However, no clear pattern was discernable for any of the components
(see Appendix 6 for results and list of species and associated dietary quality
values). Comparisons between sites indicated that the Azima tetracantha protein
value was lower in Shamwari as compared to the other reserves. The Schotia
afra value also was found to be lower in Kariega than in the AMC. The NDF
value of Acacia karroo was found to be lower in Kariega and Shamwari than
Blaauwbosch. Cynodon dactylon also had a lower NDF value in Asante Sana as
compared to the AMC and NCA. Generally values of protein, phosphorus and
NDF were found to be fairly consistent across sites (see Appendix 6 for dietary
quality values). The comparisons of the PDI’s of the AMC and NCA revealed no
significant differences between the two sites for protein, phosphorus or NDF (see
Appendix 6 for results).
80
As plant dietary quality measures were only based on single sample values for
each species per season within a reserve, and the lack of any discernable
pattern or significant statistical relationship, these values were not considered for
the explanation of the observed body condition scores. Although the ANOVA of
faecal phosphorus indicated a significant difference in phosphorus between
seasons and sites, due to its lack of contribution to the observed body conditions
as shown by the multiple regression, it was excluded from explanations of
observed body condition scores.
4.5 Discussion
4.5.1 Population specific patterns
Energetic and nutrient requirements within life stages are assumed to be
constant across locations as they are based on elephant life history and
physiology, and therefore any observed differences in body condition must be
attributable to some site-specific variables affecting condition.
The observed lower condition estimates in the AMC (Figs. 4.1, 4.3 – 4.8) for all
seasons and both energy classes (with the exception of non-energy stressed
individuals in winter) were correlated with an exceptionally high density
compared to the other reserves, as well as low rainfall during the study period
(see Table 4.1). This supports my prediction that poorer condition would be
associated with areas of high elephant density and low rainfall (i.e. productivity).
Comparisons with dietary quality measures revealed that, with the exception of
summer, these lower body condition scores were also associated with generally
lower faecal protein and higher faecal NDF values as compared to other reserves
(Tables 4.2 & 4.6), supporting my prediction that lower conditions estimates
would be associated with areas of lower dietary quality. If the lower estimate of
protein for maintenance requirements (6%, Malpas 1977) is used, this data
suggests that elephants in the AMC are on a sub-maintenance diet for nine
months of the year as compared to three months in the NCA, Asante Sana and
Blaauwbosch, with elephants in Shamwari and Kariega receiving sufficient
81
protein throughout the year. Although the estimate by Malpas (1977) refers to the
dietary protein requirements, and is thus as such not directly comparable to the
faecal protein values obtained, the inclusion of a high proportion of browse in the
diet would result in an over- and not underestimation of dietary protein, due to the
higher levels of protein-precipitating secondary compounds present in browse
(Mould & Robbins 1981; Irwin et al. 1993). Elephants in the AMC thus appear to
be relying on short flush periods after good rains to boost their protein intake and
condition, which then deteriorates during the rest of the year. This suggests that
they would be particularly vulnerable to becoming protein limited if an extended
period of drought were to occur.
As predicted, the slightly better condition exhibited by energy stressed elephants

in Asante Sana in spring (Fig. 4.3) was associated with a much higher than
average rainfall for the study period (Table 4.1), as well as consistently good
rains recorded from February to June 2007 (Appendix 7). This was not reflected
in the dietary quality measures. These higher body conditions scores may,
however, be related to the relative foraging effort at that time. Even if the
increased rainfall did not affect the quality of the available forage, the abundance
of vegetation would increase (Coe et al. 1976), thereby providing the elephants
with more food. Additionally, increased rainfall may lead to the formation of
smaller puddles and pools, thereby eliminating the need to travel larger distances
to and from water sources. This decrease in daily activity would contribute to a
decrease energy demands, thus allowing more energy to be located to storage,
and ultimately contributing to higher body condition scores.
Although the better condition exhibited by energy stressed individuals in summer

(Fig. 4.5) was associated with a large increase in protein and a drop in NDF
values at that time (Tables 4.2 & 4.6), the slightly better conditions exhibited in
Blaauwbosch by non-energy stressed individuals in spring (Fig. 4.4) was not
supported by dietary quality measures. The better condition exhibited during the
spring/summer period was also contrary to my expectations based on rainfall.
82
Based on the low rainfall in this area, it would be expected that these elephants
would exhibit lower condition estimates than some of the other reserves. This is
especially true for the spring/summer sampling period, as it was preceded by no
rains from June to September 2007 (Appendix 7). Although the improved
condition of elephants in spring remains unexplained, the increase in faecal
protein and decrease in faecal NDF during the summer period is associated with
the fruiting of the prickly pear, Opuntia ficus-indica (Brutsch & Zimmermann
1993), an invasive that can be found in large numbers throughout the reserve
(pers. obs.). Elephants were seen to feed extensively on these fruits (pers. obs.),
thus accounting for the higher dietary quality values and better body conditions
observed.
The slightly lower body condition of energy stressed individuals in the NCA in
summer (Fig. 4.5) and autumn (Fig. 4.7) were supported by higher values of
faecal NDF (Table 4.6) and lower values of faecal protein (Table 4.2) than other
reserves during this time, although faecal protein values were only at sub-
maintenance level (6%; Malpas 1977) during autumn. The slightly lower body
condition estimates obtained for the NCA elephants are also supported by the
much higher elephant density than all but the AMC population, again highlighting
this as a driver of the observed body condition scores.
These results support those obtained from the regression model, both for dietary
quality and for the site specific drivers (Appendix 4), which highlighted elephant
density, rainfall during the study period, faecal protein and faecal NDF as drivers
of the observed elephant body conditions. Although not all relationships were
significant, there was a general trend of populations with large proportions of
elephants with higher body conditions (BC 5 and 6) associated with lower
elephant densities, higher rainfall during the study period, higher faecal protein
and lower faecal NDF (Figs. 4.15 – 4.18). The reverse trend was found for the
lower body conditions (BC 3 and 4), with populations with larger proportions of
elephants with body condition scores of 3 or 4 being associated with higher
83
elephant densities, lower rainfall during the study period, lower faecal protein and
higher faecal nitrogen. It is important to note that the proportion of elephants
within the four body condition scores (BC 3, 4, 5 & 6) within a population add up
to one, and that the reverse pattern observed between the two highest and two
lowest body condition scores are thus due to auto-correlation of the data, and are
both illustrating the same trend.
The graph depicting the relationship between faecal protein and the proportion of
the population within a specific body condition (Fig. 4.17) indicates that at
approximately 8.5% faecal protein, the point at which the two lower body
condition (BC 3 & 4) lines meet the X-axis (i.e. proportion = 0), none of the
population falls within the lower body condition scores. This suggests that 8.5%
faecal protein could possibly be the lower threshold value of faecal protein
needed for the maintenance of elephant body condition within a population. A
similar trend can be seen between the proportion of the population and faecal
NDF (Fig. 4.18), where none of the population falls within the two lower body
condition scores at the point where their lines (BC 3 & 4) converge, at
approximately 71% faecal NDF. This suggests that 71% faecal NDF could be the
upper threshold above which elephant body condition begins to deteriorate (i.e.
some of the population are in poor condition). These observations, however,
need to be further tested using a larger dataset, and provide a useful working
hypothesis.
All four regression graphs (Figs. 4.15 – 4.18) show two clusters of data, with the
AMC clustered away from the other populations. It is thus possible that the
correlations between the data could simply be due to Simpson’s paradox.
However, there is no evidence that Addo is a separate population of data, and I
feel that the AMC is only the extreme of a gradient, and that the processes
determining elephant body condition are the same within all the reserves. I am
thus confident that these regressions are valid. However, the model fit R2 was
quite low for both models (site specific drivers R2 = 0.354; dietary quality R2 =
84
0.274), and thus only explained 35.4% and 27.4%, respectively, of the variability
observed, and it is therefore possible that some other, un-measured parameter
could have contributed to the observed body condition distributions.
The observed variation in body condition of elephants across seasons in both

energy classes in the AMC (Fig. 4.9) seemed to be roughly correlated with the
rainfall recorded three months prior to the sampling dates (Appendix 7). Good
body conditions in June were related to the peak in rainfall recorded in March,
April and May, and the decrease in condition recorded in September was related
to the decrease in rainfall recorded in June, July and August. Rainfall remained
fairly constant, with no change recorded in condition in December. However,
although good rains were recorded for December/January, body condition
decreased from summer to autumn. With the exception of the good body
condition scores observed in winter, body conditions were correlated with dietary
quality measures (Tables 4.4 & 4.6). This could be seen especially in the large
drop in condition from winter to spring, which was associated with a significant
increase in faecal NDF and a significant decrease in faecal protein at that time.
Although the changes were not always significant, elephants of the NCA showed
a similar trend in body condition (Fig. 4.10) in response to rainfall (Appendix 7),
as illustrated by the AMC elephants. However, whereas the AMC elephants
decreased in condition from summer to autumn, elephants from the NCA
remained in similar condition. Comparisons with dietary quality measures
revealed no correlations between the observed conditions and dietary quality.
The lack of variation in body condition scores for any of the energy classes
across seasons in Kariega (Fig. 4.1) could be attributed to the high rainfall
experienced almost throughout the year (Appendix 7), as well as the relatively
large size of the reserve and low elephant density. Although the elephants were
not in exceptional condition, their consistent condition throughout the course of
the year suggests that they were receiving sufficient nutrients for maintenance.
85
This is supported by the consistently good faecal protein values obtained in the
reserve (Table 4.4).
The drop in body condition score experienced by energy stressed elephants in

Asante Sana in summer (Fig. 4.12) could not be attributed to rainfall. Although no
rainfall was recorded for September, almost 90 mm of rain was recorded in
October (Appendix 7), and rainfall in the reserve was much higher during the
study period than the annual average. Dietary quality measures showed a mixed
response in summer, with both NDF and protein values increasing (Tables 4.4 &
4.6), and it is therefore uncertain whether the observed changes in body
condition could be attributed to changes in dietary quality. However, elephants in
Asante Sana maintained good body conditions for most part of the year, possibly
due to their very low density.
The lack of variation in body condition of elephants at Blaauwbosch (Fig. 4.13)

was contrary to what was expected, not only based on the low average annual
rainfall and small reserve size, but also the absence of rainfall from June to
September 2007 (as discussed above). With the exception of higher body
condition estimates for non-energy stressed individuals being associated with a
drop in NDF during spring (Table 4.6), dietary quality measures were not
correlated with the body conditions observed. The fairly consistent conditions
were, however, supported by the low elephant density.
With the exception of energy stressed elephants in summer, Shamwari elephants

maintained condition throughout the year (Fig. 4.14). This is in accordance with
expectations, as Shamwari has a relatively low population density and rain falls
throughout the year (Appendix 7). The decrease in condition experienced in
summer could be explained by the two low rainfall months in September and
October 2007. Dietary quality remained high throughout the study period (Tables
4.4 & 4.6), thus supporting the good body conditions observed.
86
Energy stressed and non-energy stressed classes across seasons for Kariega,
Asante Sana, Blaauwbosch and Shamwari (Figs. 4.11 – 4.14) showed an
inconsistency in their trends, with energy stressed individuals sometimes in
better and sometimes in worse condition than non-energy stressed individuals,
and this pattern being different for each site. It is important to note that the
differences in condition between these groups were fairly small and non-
significant, and that these patterns could have been created by small, natural
variations in condition of the individual elephants comprising the classes over
time.
4.5.3 Dietary quality variation

With the exception of Asante Sana and the NCA, dietary quality followed the
rainfall pattern observed at each site. Body condition on the other hand seemed
to be correlated with the rainfall recorded roughly three months before recorded
body conditions. Various authors have reported responses in forage resources to
changes in rainfall (e.g. Sinclair 1974; Coe et al. 1976; Phillipson 1975; Owen-
Smith 1990), with values of nitrogen positively correlated with rainfall (McCullagh
1969b; Sinclair 1974; Erasmus et al. 1978; Hall-Martin et al. 1982; Van der Waal
et al. 2003), and values of fibre negatively correlated with rainfall (Hall-Martin et
al. 1982). Rutherford (1984) showed that browse and grass/forb species had
different responses to rainfall. Whereas grasses and forbs responded rapidly to
changes in rainfall, browse species were slower to respond, generally responding
to the previous season’s rainfall. This was supported by Hoffman & Cowling
(1990), who showed that grasses of the eastern Karroo showed marked
responses to seasonal and short term rainfall, whereas shrubs of the sub-tropical
thickets had much less dramatic responses to rainfall. As herbivores select for
the forage item highest in nutrition based on availability (Owen-Smith 2002), it
would be expected that they would selectively feed on the flush of rapidly
growing, nutritious grasses and forbs directly after rains, thus explaining the
correlation between rainfall and increased dietary quality. This is supported by
the elephant dietary study of Davis (2005), who showed that herbaceous plants
87
became the dominant component in the diet during summer, which was also the
period with the highest recorded rainfall. The delayed response in elephant body
condition could be a result of the slower response of browse species to rainfall.
Browse has been shown to contribute between 39 – 54% of the elephant diet in
succulent thicket (Davis 2005; Wolmarans 2006; Landman et al. 2008), and as
such greatly contributes to the nutrition and condition of elephants. Alternatively,
it could represent a delay in the response of body condition to increased dietary
quality (Malpas 1977).
As herbivores are ultimately limited by their food resources (Sinclair 1974, 1975;
Skogland 1983; Mduma et al. 1999), their body condition is linked to the quality
and quantity of available forage (Price & White 1985; Brochus et al. 1988;
Choquenot 1991; Pettorelli et al. 2005). Numerous studies have demonstrated a
decrease in overall condition of herbivores in areas of high density due to a
decrease in the availability of forage (Ruffner & Carothers 1982; Choquenot 1991;
Swihart et al. 1998; Mduma et al. 1999; Van der Waal et al. 2003; Keyser et al.
2005; Stewart et al. 2005). Rainfall also directly affects primary productivity, with
a decrease in rainfall being associated with a decrease in forage availability
(Sinclair 1974; Coe et al. 1976; Phillipson 1975; Owen-Smith 1990; Mduma et al.
1999). This is supported by my findings, which illustrated a correlation between
dietary quality (i.e. faecal protein and faecal NDF) and rainfall, as well as a
decrease in elephant body condition with increasing elephant density, and
emphasises these factors as drivers of body condition through their effect on
resource availability in this landscape. This supports my hypothesis that elephant
body condition will vary as a function of primary productivity, with lower primary
productivity (due to rainfall and elephant density) being associated with lower
body condition scores.
My study has shown that, for elephant populations occurring in the Eastern Cape,
variables that affect condition cannot be considered in isolation, as they interact
with one another in different ways. It is this interaction between rainfall, elephant
88
density and forage quality that ultimately drives the observed variations in body
condition over space and time.
Although variation in dietary quality and rainfall was observed within all reserves,
the much more dramatic response in body condition observed in the AMC, as
compared to other reserves, demonstrates that the high density in the AMC has
resulted in the decrease in available resources in this reserve (as shown by
Penzhorn et al. 1974, Barratt & Hall-Martin 1991, Lombard et al. 2001), causing
an increase in competition for the limited resources. However, the individual
responses are not yet shown at the population level, as is evident by the lack of
density dependent effects on population performance (Gough & Kerley 2006). A
study by McGregor & Butler (2008) found that mortality due to hypothermia in
Angora goats was related to body condition, with a significantly increase in
mortality rates experienced when body conditions decreased below a specific
point. This was found to be confounded by goat density, with areas of high goat
density experiencing higher mortality rates (McGregor & Butler 2008). This
suggests that, although body conditions in the AMC are currently not low enough
to induce density dependence, a prolonged period of inadequate food supply (i.e.
drought) may result in an increased mortality rate, which would be exacerbated
by the high elephant density within the park. The findings of McGregor & Butler
(2008) that show that increases in mortality occur when a threshold is crossed
suggests that, when such mortalities are observed, the resources will already
have been overutilized. This concurs with Gough & Kerley’s (2006) postulated
catastrophic expression of density dependence through a collapse of resources
in the AMC, rather than a gradual increase in mortality and decline in fecundity.
With the exception of one or two observations, the lack of any significant
difference between energy stressed and non-energy stressed individuals in all
reserves but the AMC indicate that, in these reserves, the increased energy
requirements of these life stages (i.e. lactation, old age and weaning) are met by
sufficient quantity and quality of forage. This is in contrast to the AMC, where not
89
only energy stressed, but also non-energy stressed individuals are in poorer
condition than other reserves, and again highlights the fact that the high density
over time of this population has already negatively affected their resources.
The above results thus support the hypotheses that 1) elephant body condition
will vary across sites as a function of primary productivity, and 2) that elephant
body condition will vary across sites as a function of dietary quality.
90
CHAPTER 5
CONCLUDING DISCUSSION
¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯¯
5.1 Body condition scores
The need for an alternative, non-invasive method of determining condition for the
study of wild populations has given rise to the use of visual body condition
estimations. Although not yet as widely used as the more classic measures such
as Kidney Fat Index or Bone Marrow Fat, studies have shown that they can be
used to give an accurate reflection of nutritional state of the animal at the time of
observation (Riney 1960; Poole 1989; Grant et al. 1995; Wemmer et al. 2006).
This has been supported by my study, which has shown the visual estimation of
body condition to be a inexpensive, non-invasive, reliable and repeatable
measure for the determination of condition in African elephants, and which allows
for comparisons between seasons and sites and over the life history of the
animals.
5.2 Responses in body condition

The observed responses in body condition of the various life stages were in
accordance with my predictions. The lower condition estimates of energy
stressed individuals within the AMC illustrates that these life stages are more
sensitive to resource limitation, an observation which is supported by findings
from other authors (Corfield 1973; Sharp 1982; Clutton-Brock 1984; Tyler 1986;
Freeland & Choquenot 1990). This suggests that these life stages should be the
focus of any study which aims to detect resource limitation in a population, as
they will be the first to respond when resources become limiting.
The lack of response in weaned individuals did not follow my predictions.

According to Fowler (1987) and Eberhardt (2002), long-lived vertebrate vital rates
have different sensitivities to resource limitation, with juvenile mortality being the
most sensitive and therefore the first to respond to a decrease in resource
91
availability. This was supported by findings of Corfield (1973) on the Tsavo
elephant die-off. Although Corfield (1973) referred to juveniles as elephants up to
the age of four, it is thought that other author’s generally refer to the survival over
the first year of life, thereby highlighting the sensitivity of only this initial, short
period. My findings suggest that weaned juveniles are not experiencing any
increased nutritional stress due to weaning, probably due to the long, slow
weaning process exhibited by elephants (Langer 2003). Additionally, the drought
resistant nature of the thicket (Stuart-Hill & Aucamp 1993) dominating most of the
parks studied may be contributing to this lack of response. This suggests that this
particular life history is not that important for the determination of resource
limitation within populations here. These findings are supported by the low
juvenile mortality rate found in the AMC by Gough & Kerley (2006).
The lower body condition estimates obtained for lactating females, on the other
hand, highlights the energy demands of this stage of reproduction (Sharp 1982;
Gittleman & Thompson 1988; Clutton-Brock et al. 1989; Randolph & Cameron
2001). This suggests even more serious energy implications for cows that are
both pregnant and lactating. One evolutionary response to maximize fitness
would be to abort or abandon calves in times of severe nutritional stress (Laws
1969; Wasser & Barash 1983; Mellado et al. 2004), allowing the mother to
survive the stress and possibly breed again at a later stage. Evidence for
abortion in stressed populations of elephants would thus be valuable, and the
availability of new methods for monitoring pregnancy in individuals (e.g. faecal
steroid tests, Stoops et al. 1999; Foley et al. 2000; Ostrowski et al. 2005) would
allow this hypothesis to be tested.
Another response to a decrease in resource availability would be to cease

breeding until environmental conditions become favourable again (Stewart et al.
2005), or synchronize breeding with resource availability (Wittemyer et al. 2007a).
This would increase the chance of calf survival, thereby maximizing the female’s
reproductive success, and ultimately fitness (Wasser & Barash 1983). Although
92
some evidence of a positive relationship between breeding and rainfall (and thus
primary productivity) has been found in the AMC (Gough & Kerley 2006),
population growth rate is not showing any sign of slowing down, despite an
obvious decline in resource availability (Gough & Kerley 2006). Seydack &
Bigalke (1992) suggested that such a response may be due to the ratio of so-
called production to maintenance nutrients found in the eastern Cape. They
found that populations of bushpigs exhibited regional differences in their relative
investment in reproduction and maintenance, and that this was related to nutrient
availability and its effect on metabolic turnover rates. Specifically, they found that
bushpig populations in the eastern Cape had higher reproductive investments
than bushpigs in the southern Cape, who in turn had higher somatic investments.
This was also found to be true for elephant populations in these areas (Seydack
et al. 2000). Individuals in the eastern Cape populations would thus have an
adaptive strategy of maximizing current reproduction even at the risk of reduced
survival (Seydack & Bigalke 1992). This, along with the buffering effect of slow
demography (Owen-Smith 1988), could explain the high elephant population
growth rate in the AMC, despite a decline in condition and decreasing resource
availability. Along with loss of tooth function, the very poor condition exhibited by
old females in the AMC could be due to this higher investment in reproduction,
resulting in even further reduced condition, and thus fitness, at this late life stage.
My findings suggest that, despite no current evidence of density dependence

(Gough & Kerley 2006), the AMC elephant population are experiencing resource
limitation as shown by their body condition. As large, long-lived herbivores are
buffered against short-term food insufficiency (Owen-Smith 1988), they should be
slower to respond to declining resources. As indicated by McGregor & Butler
(2008) for goats, there may be some critical body condition at which elephants
become susceptible to environmental stressors, with subsequent effects on
survival. It is not clear, however, what body condition would be the threshold at
which this will occur.
93
My findings also suggest that, based on observed variations in both dietary
quality and elephant body condition, thicket is not as stable in its provision of
nutrients as previously thought (Everard 1987; Stuart-Hill & Aucamp 1993), even
for elephants who are not really considered to be that vulnerable to changes in
resources (Demment & Van Soest 1985; Owen-Smith 1988; Duncan et al. 1990).
Additionally, my regressions (Figs 4.17 & 4.18) highlighted values of faecal
protein (c. 8.5%) and faecal NDF (c. 71%), below which body condition appeared
to deteriorate. This suggests that it may be possible to establish threshold values
of dietary quality that may serve as a benchmark for the determination of the
condition of an elephant population.
This study set out to investigate the effect of life stage and seasonal and site-
specific differences in productivity on elephant body condition. My results have
indicated that elephant body condition does vary between life stages, with energy
stressed individuals, and in particular, lactating and old females, exhibiting lower
body conditions than non-energy stressed individuals. I have also shown that
elephant body condition responds to seasonal and site-specific changes in
productivity, with these differences in condition across space and time being
attributed to variations in elephant density, rainfall and associated faecal protein
and NDF. Specifically, I have shown that elephants exhibit lower body conditions
at sites and during seasons with lower primary productivity (as based on rainfall
and dietary quality).
5.3 Future Research

The use of different indices of condition found within the literature make it difficult
to compare results obtained from various studies. This is especially true for
comparisons between visual condition indices and biological condition indices
(such as Kidney Fat Index or Bone Marrow Fat). It is suggest that, if culling of
elephants were ever to take place, it would be beneficial to look at these more
traditional body condition measures and calibrate these with the observed visual
body condition estimates. Alternatively, the use of either catabolic breakdown
94
products in the urine, such as creatinine, or faecal stress hormones (Foley et al.
2000) could be used to benchmark visual body condition scores. This would
allow for possible refinement and greater degree of confidence in the use of
visual body condition estimates, and would allow for better comparisons of
condition obtained via different methods.
One shortcoming of this study was the lack of data on new born calves (i.e.
under one year of age), as well as females who were both pregnant and lactating.
It is suggested that future research include these two categories, as it is
expected that they would be the even more affected by nutritional stress.
Additionally, the lack of seasonal replication makes it difficult to track long-term
responses and natural variations in condition, and it is suggested that future
research follow changes in elephant body condition of a population for at least
three years to provide a minimal seasonal replication.
Although my study showed responses in body condition to changes in rainfall, it

is suggested that future research measure body condition across a broader
landscape with larger differences in soil nutrients and rainfall. This will allow a
greater insight into the sensitivity of elephant condition to changes in these
parameters. I also suggest that future research compare body conditions of
elephants occurring in different biomes, which will give an indication of the
responses of elephant condition in various landscapes. Additionally, I suggest
that the rate of recovery of body condition should be measured in a system
where distinct wet and dry seasons are experienced. This will aid in our
understanding of the possible time-lags between observed condition and
resource availability.
Lastly, I also suggest that it may be beneficial to further investigate the possible
link between critical faecal protein and NDF values and body condition, by using
a larger dataset of elephant body condition scores and faecal samples to verify
the suggested dietary quality thresholds found in my data. If such threshold
95
values can be determined it could be possible to predict the body conditions of
elephants in a population by the collection and analysis of faecal samples. This
would be especially useful for the determination of condition of elephant
populations found in the forests of West Africa, which are often hard to observe
and study (Merz 1986; Eggert et al. 2003).
96
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Appendix 1: Samples sizes obtained for the various sites,
seasons and life stages.
Table A1.1 Sample sizes obtained for the Addo Main Camp for comparisons
between life stage and seasons.
Season
Life‐history Winter Spring Summer Autumn
Old 5 4 4 4
Lactating 33 36 38 53
Sub‐adult/non‐
lactating 7 8 12 15
Weaning 11 14 18 50
Table A1.2 Sample sizes obtained for all sites for comparisons between sites,
seasons and energy classes. ES = Energy stressed, NES = non-energy stressed,
n/s = not sampled.
Season
Site Winter Spring Summer Autumn
ES NES ES NES ES NES ES NES
Addo Main Camp 49 7 54 8 60 12 107 15
Nyathi Concession Area 20 6 18 6 20 13 20 12
Asante Sana 4 5 4 5 4 5 4 5
Shamwari 16 19 n/s n/s 21 19 18 20
Kariega 7 8 7 6 8 6 7 8
Blaauwbosch 6 4 6 4 6 4 6 4
111
Appendix 2: Results of log-linear analysis of body condition and
life stage in the AMC.
Table A2.1 Log-linear results for all k-factor interactions. Significant p values in
bold and italicized (Statistica 8.0; StatSoft Inc. 2007).
Results of Fitting all K-Factor Interactions

These are simultaneous tests that all K-Factor
Interactions are simultaneously Zero.
Degrs.of Max.Lik. Probab. Pearson Probab.
K-Factor Freedom Chi-squ. p Chi-squ p
1 9 278.4039 0.000000 372.0410 0.000000
2 27 111.9832 0.000000 114.9325 0.000000
3 27 17.3107 0.923177 18.7457 0.879081
Table A2.2 Log-linear results for all marginal and partial associations. Factor 1 =
season, 2 = life stage, 3 = body condition category. Significant interactions in
Tests of Marginal and Partial Association (Life-history x season x BC)

Degrs.of Prt.Ass. Prt.Ass. Mrg.Ass. Mrg.Ass.
Effect Freedom Chi-sqr. p Chi-sqr. p
1 3 27.5852 0.000004 27.5852 0.000004
2 3 138.7952 0.000000 138.7952 0.000000
3 3 112.0238 0.000000 112.0238 0.000000
12 9 25.6307 0.002347 14.0929 0.119060
13 9 57.6339 0.000000 46.0959 0.000001
23 9 51.7944 0.000000 40.2565 0.000007
112
Appendix 3: Results of log-linear analysis of body condition for
all sites and seasons.
Table A3.1 Log-linear results for all k-factor interactions. Significant p values in
Results of Fitting all K-Factor Interactions

These are simultaneous tests that all K-Factor
Interactions are simultaneously Zero.
Degrs.of Max.Lik. Probab. Pearson Probab.
K-Factor Freedom Chi-squ. p Chi-squ p
1 12 607.7183 0.000000 1059.672 0.000000
2 50 348.9712 0.000000 359.766 0.000000
3 84 73.0085 0.798441 79.197 0.627849
4 45 28.0507 0.977549 27.378 0.982316
Table A3.2 Log-linear results for all marginal and partial associations. Factor 1 =
site, 2 = season, 3 = energy class, 4 = body condition category. Significant
interactions in bold and italicized (Statistica 8.0; StatSoft Inc. 2007).
Tests of Marginal and Partial Association (site x season x energy class x BC)
Degrs.of Prt.Ass. Prt.Ass. Mrg.Ass. Mrg.Ass.
Effect Freedom Chi-sqr. p Chi-sqr. p
1 5 349.1582 0.000000 349.1582 0.000000
2 3 32.3711 0.000000 32.3711 0.000000
3 1 37.4452 0.000000 37.4452 0.000000
4 3 188.7439 0.000000 188.7439 0.000000
12 15 39.1643 0.000606 46.4120 0.000046
13 5 11.7862 0.037838 25.6236 0.000106
14 15 184.6691 0.000000 206.6986 0.000000
23 3 9.7569 0.020749 5.4637 0.140823
24 9 50.5459 0.000000 54.4457 0.000000
34 3 16.0537 0.001106 26.5436 0.000007
123 15 9.1575 0.869138 7.5602 0.940202
124 45 44.6454 0.486873 41.8600 0.605747
134 15 13.6190 0.554597 11.0921 0.746041
234 9 5.3741 0.800555 9.6937 0.375844
113
Appendix 4: Results of multinomial regression models
Table A4.1 The full and reduced proportional odds ordinal logistic regression
model for body condition and site specific drivers. Significant p values bold and
italicized (Design; Harrell 2008).
Full model has an R2 of 0.360
Coef S.E. Wald Z P

Size -0.00515 0.008101 -0.64 0.5248
Density -0.92332 0.267565 -3.45 0.0006
Long term
rainfall -0.00075 0.003459 -0.22 0.828
Study period
rainfall 0.002078 0.001021 2.03 0.0419
Years 0.014412 0.019681 0.73 0.464
Reduced model has an R2 of 0.354
Coef S.E. Wald Z P

Density -0.704 0.056769 -12.4 <0.0005
Study period
rainfall 0.000941 0.000816 1.15 0.2493
Table A4.2 The full and reduced proportional odds ordinal logistic regression
model for body condition and dietary quality. Significant p values italicized
(Design; Harrell 2008).
Full model has an R2 of 0.275
Coef S.E. Wald Z P

Protein -1.0639 0.4755 -2.24 0.0252
P 7.5473 6.5302 1.16 0.2478
NDF -0.7744 0.1142 -6.78 <0.0005
Reduced model has an R2 of 0.274
Coef S.E. Wald Z P

Protein -0.5845 0.2239 -2.61 0.009
NDF -0.7274 0.1035 -7.03 <0.0005
114
Appendix 5: Results of ANOVAs for faecal dietary quality.
Table A5.1 Results of ANOVAs for protein, phosphorus and NDF for all sites and
seasons (Statistica 8.0; StatSoft Inc. 2007).
Univariate Tests of Significance for Value (protein)

Sigma-restricted parameterization
Effective hypothesis decomposition
SS Degr. of MS F p
Effect Freedom
Intercept 0
Site 20.1762 4 5.04404 2.97576 0.023259
Season 116.6587 2 58.32934 34.41176 0.000000
Site*Season 67.5561 14 4.82543 2.84680 0.001384
Error 155.9437 92 1.69504
Univariate Tests of Significance for Log Value (Phosphorus)
SS Degr. of MS F p
Effect Freedom
Intercept 0
Site 0.198517 4 0.049629 6.40346 0.000137
Season 0.408720 2 0.204360 26.36775 0.000000
Site*Season 0.338406 14 0.024172 3.11880 0.000528
Error 0.713034 92 0.007750
Univariate Tests of Significance for Value (NDF)
SS Degr. of MS F p
Effect Freedom
Intercept 0
Site 185.621 4 46.40520 3.821099 0.006450
Season 173.553 2 86.77640 7.145346 0.001311
Site*Season 352.534 14 25.18097 2.073452 0.020469
Error 1105.146 91 12.14446
115
Appendix 6: Plant dietary quality (protein, phosphorus and NDF)
values and results of overlapping plant species and PDI’s.
Table A6.1 Overlapping dietary plant species with corresponding dietary quality
measures for summer and winter. Site names: A = AMC, N = NCA, K = Kariega,
B = Blaauwbosch, S = Shamwari, AS = Asante Sana.
SPECIES SITE SEASON PROTEIN P NDF

Azima tetracantha Summer 15.9 0.16 37.5
A
Winter 15.4 0.48 33.8
Summer 13.9 0.19 32.2
N
Winter 16.5 0.15 40.1
Summer 10.4 0.12 41.3
K
Winter 12.1 0.13 27.6
Summer 15.6 0.15 29.5
B
Winter 12.2 0.13 42.4
Summer 8.1 0.14 49.1
S
Winter 9.0 0.13 35.0
Schotia afra Summer 11.3 0.19 48.7
A
Winter 9.6 0.13 54.7
Summer 8.9 0.14 54.1
N
Winter 15.4 0.11 51.1
Summer 6.4 0.10 64.8
K
Winter 7.1 0.11 50.5
Cynodon dactylon Summer 10.3 0.36 64.9
A
Winter 12.9 0.22 68.5
Summer 7.9 0.31 68.3
N
Winter 12.1 0.26 67.5
Summer 16.0 0.48 61.1
AS
Winter 7.0 0.24 66.7
Acacia karroo Summer 9.3 0.11 38.5
K
Winter 9.8 0.11 22.7
Summer 14.9 0.15 34.0
AS
Winter 8.4 0.13 40.8
Summer 8.4 0.13 48.0
B
Winter 8.0 0.16 59.4
Summer 14.4 0.14 32.1
S
Winter 9.3 0.13 48.9
116
Table A6.2 Results of paired t-tests for protein, phosphorus and NDF values of
overlapping dietary plant species (those species occurring in the diet on three or
more sites) (Statistica 8.0; StatSoft Inc. 2007).
MEASURE SPECIES N df p
PROTEIN Azima
0.793716
tetracantha 5 4
Schotia afra 3 2 0.529607
Cynodon
0.880700
dactylon 3 2
Acacia karroo 4 3 0.191117
P Azima
0.483955
tetracantha 5 4
Cynodon
0.120616
dactylon 3 2
NDF Azima
0.718255
tetracantha 5 4
Cynodon
0.280318
dactylon 3 2
117
Table A6.3 Results of ANOVAs for protein, phosphorus and NDF values of the
PDI’s of the AMC and NCA (Statistica 8.0; StatSoft Inc. 2007).
Univariate Tests of Significance for Value (protein)

SS Degr. of MS F p
Effect Freedom
Intercept 4850.650 1 4850.650 410.2148 0.000000
Site 21.486 1 21.486 1.8171 0.183991
Season 26.871 3 8.957 0.7575 0.523496
Site*Season 2.639 3 0.880 0.0744 0.973461
Error 567.584 48 11.825
Univariate Tests of Significance for Value (Phosphorus)
SS Degr. of MS F p
Effect Freedom
Intercept 1.666350 1 1.666350 169.9119 0.000000
Site 0.000457 1 0.000457 0.0466 0.829980
Season 0.025907 3 0.008636 0.8806 0.457846
Site*Season 0.033543 3 0.011181 1.1401 0.342415
Error 0.470743 48 0.009807
Univariate Tests of Significance for Value (NDF)
SS Degr. of MS F p
Effect Freedom
Intercept 149181.8 1 149181.8 558.1263 0.000000
Site 763.2 1 763.2 2.8555 0.097549
Season 235.7 3 78.6 0.2939 0.829609
Site*Season 453.1 3 151.0 0.5650 0.640722
Error 12829.9 48 267.3
118
Appendix 7: Rainfall recorded at each site for 2007/2008
Fig. A7.1 Rainfall recorded for Asante Sana from January 2007 to April 2008.
Long term mean 360 mm.
Fig. A7.2 Rainfall recorded for Blaauwbosch from January 2007 to April 2008.
Long term mean 300 mm.
119
Fig. A7.3 Rainfall recorded for Shamwari from January 2007 to April 2008. Long
term mean 550 mm.
Fig. A7.4 Rainfall recorded for Kariega from January 2007 to April 2008. Long
term mean 640 mm.
120
Fig. A7.5 Rainfall recorded for the AMC and NCA from January 2007 to April
2008. Long term mean 396 mm.
121
122

Detecting Changes in Elephant Body Condition in Relation To

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Detecting Changes in Elephant Body Condition in Relation To

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Detecting changes in elephant body

Supervisor: Prof G.I.H. Kerley

1.2 The elephant problem

In addition to their effects on plant communities, elephants also impact individual

The degree to which elephants impact their environment is a function of their

1.3 Elephant life history and population regulation

Large mammal population regulation generally occurs via one of two

1.4 Caughley’s framework

Fig. 1.1 A graphical representation of Caughley’s interferential plant-herbivore

1.5 Body condition as indicator of response

1.6 Elephants as herbivores

1.7 Measuring diet quality

1.8 The elephant problem in the Eastern Cape

1.9 Rationale, objectives and research approach

1. Determine whether energetically constrained elephant life stages (i.e.

3. Determine how site differences in productivity and elephant density affect

2.2.2 Decline of the elephant

2.2.3 Current populations and distribution

2.2 Study sites and populations

2.2.1.4 Elephant population history

2.2.1.5 Elephant diet

2.2.2.5 Elephant diet

2.2.3 Asante Sana Private Game Reserve

The mean annual temperature ranges between 15 and 17.5°C, although

2.2.3.4 Elephant population history

2.2.3.5 Elephant diet

2.2.4 Shamwari Private Game Reserve

According to O’Brien (2004) the thicket is characterized by a dense tangle of

2.2.4.4 Elephant population history

2.2.4.5 Elephant diet

2.2.5 Kariega Private Game Reserve

Characteristic species include Cassine aethiopica, Euphorbia triangularis, E.

2.2.5.4 Elephant population history

2.2.5.5 Elephant diet

2.2.6.4 Elephant population history

2.2.6.5 Elephant diet

3.1.2 Body condition over lifetime

3.1.3 Condition measures

3.2 Hypotheses and aims

This will enable me to determine whether energetically constrained elephant life

Additionally, by comparing the body condition of individuals (within their

3.3 Materials and methods

3.3.2 Data collection

Where possible, photographs were taken to aid in identification of individuals, as

Graphical exploration of these results revealed that, in winter, lactating

WOld WLact WWean Wsub/non

SpOld SpLact SpWean Spsub/non

SuOld SuLact SuWean Susub/non

AuOld AuLact AuWean Ausub/non

3.5.1 Condition and life stage

3.5.2 Condition and season

Seasonal comparisons revealed that all classes except lactating females

4.1.1 Spatial variation in forage resources

4.1.2 Impact of herbivores on forage resources

The occurrence of herbivores at high densities also directly influence the

The confinement of herbivores within reserves has also created an additional

4.1.3 Effects on body condition

4.1.4 Diet quality and its assessment

The nutritional quality of ingested food is determined by its composition in terms

It is not adequate, however, to explain dietary quality only in terms of these

In addition to these requirements, herbivores also need a variety of minerals to

4.1.5 Elephant nutrient requirements