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American Journal of Research Communication www.usa-journals.com
Mwambola, et al., 2014: Vol 2(11) 102 ajrc.journal@gmail.com
Feeding preference of the African elephant (Loxodanta africana) on woody
plant species in Rubondo Island National Park (RINP), Tanzania
Simon Mwambola1*, Jasper Ijumba1, Wickson Kibasa2, Emanuel Masenga3, Ernest Eblate3
Canisius J. Kayombo4
1Nelson Mandela African Institution of Science and Technology, School of Life Sciences and
Bio-engineering, P.O Box 447, Arusha, Tanzania.
2Tanzania National Parks, Rubondo Island, P.O Box 3134, Arusha
3Tanzania Wildlife Research Institute, P.O Box 661, Arusha
4Olmotonyi Forestry Training Institute, P.0 Box 943, Arusha
*corresponding author email: mwambolas@nm-aist.ac.tz
Abstract
Studying interactions between elephant and habitats is valuable for assessing the welfare of
elephant populations and their habitats. The study was carried out in RINP between March and
July 2014 constituting wet and dry seasons to assess the feeding preference of the African
elephant (Loxodonta africana) on woody plant species. Purposive sampling was used for
gathering data from 26 sample plots, sized 50m x 50m each. Plots were designed immediately
after the herd (s) had passed on the various habitats. Frequency and percentage was used to
estimate utilization of individual plant species and their parts. Chi-square test and excel were
used to analyze feeding pattern preference. The present study found 22 species of plants in 13
families foraged by African elephants in Rubondo Island National Park. Plant species observed
to be utilized mostly by elephants included Croton sylvaticus (13.33%), Croton macrostachyus
(11.11%), Aeschynomene elaphroxylon (8.88%), Chaetacme aristata (8.88%), Ekerbegia
capensis (8.88%), Saba comorensis (6.67%), and Phoenix reclinata (6.67%). There was
difference in feeding preference between stem and leaves in woody plant species (χ2=86.462, P
< 0.05) which suggested that elephants prefer more leaves than stem. This study presents useful
information in understanding the interaction between the African elephant and vegetation in
RINP.
Keywords: African elephant (Loxodanta africana), feeding preference, island, woody plant
species.
{Citation: Simon Mwambola, Jasper Ijumba, Wickson Kibasa, Emanuel Masenga, Ernest
Eblate, Canisius J. Kayombo. Feeding preference of the African elephant (Loxodanta africana)
on woody plant species in Rubondo Island National Park (RINP), Tanzania. American Journal
of Research Communication, 2014, 2(11): 102-113} www.usa-journals.com, ISSN: 2325-4076.
Introduction
African elephants are generalist herbivores that are relatively non-selective and reliant on widely
distributed resources (Laws, 1970; Owen-Smith, 1988). ). They are mixed feeders, ingesting both
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grass and browse in varying proportions (Stephenson, 2007; Peter, 2003). Being bulk feeders
elephants prefer to spend most of their time in areas with high plant biomass (Olff et al., 2002;
Barnes, 1982). They have the ability to migrate for a long distance in searching for food, water
and when escaping hunting activities. Unlikely other connected ecosystems, Rubondo Island
National Park is a small island with no connected ecosystem. It constitute the mosaic of
different primary forest, forming dense canopy cover and lake-edge vegetation, variety of
grasslands, Acacia open woodland and forest thicket communities (TANAPA, 2003).
There is dearth of scientific published information and knowledge on the ecology of the RINP
elephant since when they were introduced between 1972 and 1972 (TANAPA, 2003; TAWIRI,
2012; TANAPA, 2014). Studying vegetation characteristics and other important resources in
conservation areas provide useful knowledge on the determinants of the density and distribution
of elephants in forests (Wing & Buss, 1970). Random stratification, systematic and purposive
approaches may be used in collection of vegetation data basing on the objectives, resources
available and nature habitats in the study area.
In unpublished report by Ernest Mjingo it was stated that, savanna eco-systems African elephants
have been observed to prefer mostly tree branches than other parts. This is partly explained by
the observed frequent pushing and falling of trees caused by elephants in savannah ecosystems
(Gadd, 2002; Mapaure & Moe, 2009). Elephant’s browsing can enhance availability of the
nutrient contents in terms of proteins and essential minerals in plant parts (Holdo, 2002). Unlike
other herbivores, elephants are adapted to use a wide range of plant species and various parts of
them (Kerley et al., 2008). Effects due to herbivory by elephants may include debarking,
breaking, felling, pushing over or uprooting young trees and seedlings during browsing, or when
young bulls engage in social displays (Smallie & O’Connor, 2000). This habit may reduce tree
species diversity, mortality and undersized growth (Chira & Kinyamario, 2009). While this
feeding behavior of elephant might be seen as disturbance to ecosystem structure, it plays an
important ecological role in savannah and forest ecosystems through maintaining suitable
habitats for numerous species (Stephenson, 2007).
Being mega herbivores, elephants need to take sufficient amount of forage and water per day.
Elephant can take up to 300kg of nutrients and 225L of water per day (Stephenson, 2007). This
means that elephant require large home range and extensive habitats to meet their basic
requirements. However in a situation where there is closed and small ecosystem like in RINP the
management of elephant population would become a great challenge when they are great in
number. The fact that RINP is an island having dry land of 236.8 Km2 with a growing elephant
population, approximately 102 individuals currently, it is important to study their feeding
behavior, to generate useful information for their management purposes. This study determined
the feeding preference of African elephants in RINP and how they interact with the forests in a
closed ecosystem.
Material and methods
Study site
Rubondo Island is located in the southwestern portion of Lake Victoria (2 o 18′ S, 31 o 50′ E) at
an altitudinal range of 1,100–1,500m above the sea level (Figure 1) (Moscovice et al. 2007).
Rainfall is bimodal. Rainy seasons take place between October to December (short) and March
to May (long), dry seasons are January and February, then June to September (TANAPA, 2003).
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Temperatures are moderate throughout the year with minimum and maximum at around 16oC
and 26oC respectively.
The habitat is dominated by mixed evergreen and semi-deciduous forest, with common species
including Croton sylvaticus, Drypetes gerrardii, and Lecaniodiscus fraxinifolius, and often with
a dense understory of lianas, or woody vines (Moscovice et al. 2007). Common native fauna
include the vervet monkey (Chlorocebus aethiops), sitatunga (Tragelaphus spekei), and
bushbuck (Tragepahalus scriptus). In addition to African elephant (Loxodonta africana) there
are 12 other introduced species on the island, including chimpanzees (Pan troglodytes), giraffes
(Giraffa camelopardalis), black and white colobus (Colobus abysinnicus), Suni antelopes
(Nestrogus moschatus) and Black rhinoceros (Diceros bicornis) (Borner, 1985). This study was
conducted between March and July, 2014. Rubondo Island was gazzeted on 18th of January 1977
and became the tenth National Park in Tanzania.
Figure 1: The map of Tanzania showing the location of Rubondo Island National Park,
Tanzania.
Sampling and Data collection
The study of woody plant species which are preferred by elephants in Rubondo Island was
conducted through plot sampling. Sample plots consisted of 50m by 50m each (Figure 2). Due to
the limited time and financial constraints plot samples were distributed purposively with
significant representative of all habitats in the park. Data collection was done either along with
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dung survey or after the herd of elephant has passed in various habitats. The main assumption
was that, elephants use almost 90% of whole habitats in the Island. Consumed plants in plot
samples were identified and unidentified species were carried to botanist in Olmotonyi Forest
Training Institute in Arusha for identification. Park rangers gave the information on encounter of
elephants either through their normal patrol or visits of elephants in ranger posts. The study was
limited by behavior of elephants of feeding during the night hours and being aggressive when
approached for close observation.
Data on feeding preference of the elephants was gathered by observing on the type of plant
species browsed, part affected, observed percentage of utilization, altitude, vegetation/ habitat
type and records of GPS coordinates of location of plot samples. Observational examination on
the elephant dung-piles was done to determine the presence of seeds on the deposited dungs.
Figure 2: Map of RINP showing distribution of vegetation sample plots.
Data analysis
Feeding preference on plant species by elephants were analyzed by computing the frequency of
foraging on individual plants and the parts affected on each species. SPSS computer software
was used to compute the Chi-square test in determining feeding preferences on parts of the
woody plant species.
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Results
Feeding preferences by elephants in Rubondo Island
Total of 26 sample plots each 50m x 50m were surveyed purposively in Rubondo Island. 20
woody plant species belonging to 13 families were found to be consumed by elephants in
Rubondo Island (Table 1). Among the identified plant species 17 were trees and 3 were woody
climbers. Fruits of Citrus limon, seeds of Phoenix reclinata and pods of Senna spectabilis were
observed in the dung piles of elephants. The lower land of the Island (close to the lake shore)
was found to be utilized more compared to the upper land.
Debarking, browsing and stem chewing were observed to be the main modes of feeding on
woody plants (Table 2). High relative frequency of utilization of encountered plant species was
encountered on; Croton sylvaticus (13.33%), Croton macrostachyus (11.11%), Aeschynomene
elaphroxylon (8.89%), Chaetacme aristata (8.89%), Ekerbegia capensis (8.89%), Saba
comorensis (6.67%), and Phoenix reclinata (6.67%) (Table 3). Observational study on degree of
destruction on individual plants found Phoenix reclinata (20-80%), Ekebergia capensis (5-80%),
Croton sylvaticus (40-60%) and Chaetacme aristata (5-30%) experienced high extent of
destruction due to foraging habit of elephants in the island (Table 2). In determination of feeding
preference on different parts of the plant we obtained that elephant in RINP prefer more leaves
than stem (χ2=86.462, P value< 0.05) (Appendix 1).
Bush buck and Vervet monkeys were observed to forage in broken Croton, Haplocoelum and
Phoenix plant species. Plants which were observed to be either broken or utilized to death
included Phoenix reclinata, Croton macrostachyus, Margaritaria discoidea, Salacia
madagascariensis and Periploca linearifolia. Terrestrial habitat is inhabited by woodland, shrub-
land, thickets and glades where by aquatic habitat is covered by marshes and Aeschynomene
elaphroxylon.
Table 1. Woody plant species encountered to be utilized by elephants in RIP, 2014
S.
No
Botanical name
Family
S.
No
Botanical name
Family
1
Haplocoelum inopleum
Sapindaceae
11
Ficus lutea
Moraceae
2
Periploca linearifolia
Asclepiadaceae
12
Ficus sycomorus
Moraceae
3
Aeschynomene
elaphroxylon
Fabaceae
13
Landolphia buchananii
Apocynaceae
4
Albizia gummifera
Fabaceae
14
Lannea fulva
Anacardiaceae
5
Annona senegalensis
Annonaceae
15
Lecaniodiscus
fraxinifolius
Sapindaceae
6
Antiaris toxicaria
Euphorbiaceae
16
Margaritaria discoidea
Euphorbiaceae
7
Chaetacme aristata
Ulmaceae
17
Phoenix reclinata
Arecaceae
8
Croton macrostachyus
Euphorbiaceae
18
Saba comorensis
Apocynaceae
9
Croton sylvaticus
Euphorbiaceae
19
Salacia madagascarensis
Celasteraceae
10
Ekebergia capensis
Meliaceae
20
Senna spectabilis
Fabaceae
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Table 2. Growth forms, parts consumption on woody plants and mode of feeding
Growth form
Part
consumed
Plant species consumed Tree Pole/sapling
Woody
climber/liana Stem Leaves Mode of feeding
Phoenix reclinata v
V v
chewing
stem/browsing
Ekebergia capensis
v
V
Debarking
Croton sylivaticus
v
V
v
Browsing
Croton macrostachyus
v
V
v
Debarking/browsing
Chaetacme aristata
v
v
Browsing
Aeschynomene
elaphroxylon
v
v Browsing
Annona senegalensis
v
v
Browsing
Lecaniodiscus fraxinifolius
v
V
Debarking
Saba comorensis
v
V
Debarking
Ficus lutea
v
V
Debarking
Periploca linearifolia
v
V
Debarking
Senna spectabilis
v
v
Browsing
Albizia gummifera
v
V
Debarking
Antiaris toxicaria
v
V
Debarking
Haplocoelum inopleum
v
v
Browsing
Landolphia buchananii
v
V
Debarking
Margaritaria discoidea
v
V
Debarking
Salacia madagascarensis
v
V
Debarking
Ficus sycomorus
v
V
Debarking
Lannea fulva
v
V
Debarking
TOTAL
16
1
3
19
9
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Table 3. Frequency of encounter rate of consumed plant species across surveyed sample
plots
S. No
Plant species
Frequency
of
encounter
Percentage
of frequency
1
Croton macrostachyus
6
13.33
2
Croton sylivaticus
5
11.11
3
Aeschynomene
elaphroxylon
4
8.89
4
Chaetacme aristata
4
8.89
5
Ekebergia capensis
4
8.89
6
Saba comerensis
3
6.67
7
Phoenix reclinata
3
6.67
8
Annona senegalensis
2
4.44
9
Haplocoelum inopleum
2
4.44
10
Lecaniodiscus fraxinifolis
2
4.44
11
Periploca linearifolia
1
2.22
12
Albiziagummifera
1
2.22
13
Antiaris toxicaria
1
2.22
14
Ficus lutea
1
2.22
15
Ficus sycomorus
1
2.22
16
Landolphia buchananii
1
2.22
17
Margaritaria discoidea
1
2.22
18
Salacia madagascarensis
1
2.22
19
Lannea fulva
1
2.22
20
Senna spectabilis
1
2.22
Discussion
Feeding preference by elephants in Rubondo Island
Out of 20 plant species identified 17 were trees, 3 woody climbers and 1 sapling. Results
suggested that there is difference in feeding preference between stem and leaves of the trees
observed, generally elephant seems to consume leaves than stem (χ2=86.462, d.f 19, P < 0.05).
Although height of the plant might be among the factor contributed to the observed difference,
but also it seems there may be a relationship between feeding preference with palatability of the
plant parts. The higher the height of stem the fewer amounts of fruits and leaves foraged by the
elephants. Our findings showed that elephants are generalist in herbivory although some extent
on preference was observed on some species ((Croton sylvaticus (13.33%), Croton
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macrostachyus (11.11%), Aeschynomene elaphroxylon (8.89%), Chaetacme aristata (8.89%),
Ekerbegia capensis (8.89%), Saba comerensis (6.67%), and Phoenix reclinata (6.67%) (Table
3). Among frequently utilized plants C. macrostachyus, P. reclinata, A. elaphroxylon, and S.
comorensis had high species richness and evenness. Availability of these plant species in various
areas makes central and northern zones to provide the suitable habitats for elephants in RINP.
Regular distribution of preferred plants close to the lake shore influences elephants to use most
of their time utilizing the habitats.
Debarking, browsing and breaking on woody plants were the common observed means of
foraging by elephants in RINP. Ecologically, being non selective elephants may influence the
change in species richness and composition. As a result decrease in species diversity may have
negative impacts to herbivores in RINP in relation to the destructive mean of feeding on
vegetation in forests including RINP. Croton macrostachyus was observed to be browsed at
stage of sapling growth form this might have been due to high concentration of nutrient such as
Potassium, Sodium, Magnesium and Phosphorus which are normally higher in young leaves than
all other leaves (Jachmann & Bell, 1985). Ficus species was categorized as low in utilization
although opportunistic surveys found that they are highly distributed along the lake shore. The
species evenness of Ekebergia capensis was found to be low but it is debarked severely almost 5
to 80%. Repeated herbivory on this plant species may lead to extinction in RINP.
Exceptional and peculiar mode of feeding was observed on consumption of P. reclinata in which
stem, leaves and fruits are foraged by elephants on various habitats. Contrary to other woody
plants species, preference of elephant to Phoenix reclinata species was on chewing the inner
section of the stem. This might have been due to easy breaking and access the juicy contents at
the inner part of stem. Leaves and fruits of Phoenix species are also well consumed by elephants
in Rubondo Island. Distribution of Phoenix reclinata is restricted along the lake shore 500m to 1
km from the water margin towards inland. Our study found that nurseries of Phoenix reclinata
were observed in the dense forests due to effect of seed dispersal by elephants. Although Phoenix
species is highly destructed and consumed by elephants causing to its death, good enough
elephants themselves disperse its seeds through foraging the fruits. Purposive and opportunistic
surveys on elephant dung piles revealed the presence of fruits of Limon citrus, pods of Senna
spectabilis and seeds of P. reclinata. Hence, elephants play important role of dispersal of plant
species in RINP. It has reported that Senna spectabilis and Limon citrus are among the more
distributed exotic plant species in the island and Senna spectabilis is likely to affect the species
composition of indigenous plants in the island (W. Kibasa per. comm., 2014). Bushbucks and
Vervet monkeys were encountered feeding on either broken or fallen trees, hence revealing the
role of elephants to other wild animals by either enhancing nutrients availability or paving the
paths for easy access to new resources.
Another exceptional observational was encountered in in aquatic habitats whereby elephants
preferred to browse on Aeschynomene elaphroxylon which is found from inland 15m to 30m in
lake water. Elephants spent not less than one hour browsing stems and leaves on Aeschynomene
elephroxylon during the night hours, probably it has high nutrient value. The results from direct
observation and analysis of food materials in the dung-piles showed that there was no change on
type of food for elephants in wet and early dry seasons Sometimes it was found that trees were
fallen or broken without foraging sign as result of either social display by sub-adult or paving the
way for easy penetration due to large body of elephants. Our general observation showed that
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showed there was no change on the type of food for elephants in wet and early dry seasons.
Almost all habitats are used by elephants with less observation in the southern part which is
inhabited by dense and spiny thickets in RINP. Similar to other tropical ecosystems vegetation in
Rubondo Island provides enough resources for the existence of this largest land living organism
in the World (Harris et al. 2008). Elephant density of 0.423 per square km2 may suggest that
there is steady environment for supporting the elephant population in the island since the forests
are still intact and healthy (Mwambola et al. 2014).
Conclusion and recommendations
Basing on our findings African elephants have successfully succeeded to adapt to survive in
forest areas since they were introduced from savanna habitats about 42 years ago. The existing
habitats are able to support the survival of elephants and other animals in the wild. The forest
structure is still closed and intact, but the rapid growth of large herbivores like elephant may lead
to over utilization of limited resources in the island. As a result forests may be transformed to
shrubs and grasslands. There is significant relationship between increased age and number of
elephants and the level of destruction in conservation areas. Hence increased number of
elephants may impose loss of suitable habitats to herbivores. Further study is needed to
understand the influence of chemical contents of the plant species on food selection by elephants
the area. This will provide foundation to explain scientifically for the causes of feeding
preference of the African elephant beyond physical observation in area which was originally
gazetted as a sanctuary for endangered species in Tanzania and Worldwide.
Acknowledgements
This study is indebted many thanks to Government of United Republic of Tanzania through
Commission for Science and Technology and The Nelson Mandela African Institution of Science
and Technology (NM-AIST) for the financial support that helped to conduct this study. Special
thanks to the management of Tanzania Wildlife Institute (TAWIRI) and Tanzania National Parks
(TANAPA) for permission to conduct research in Rubondo Island. Furthermore, Jeremiah
Johansen and Sam Ramadhan are thanked for their assistance during data gathering.
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Apendix 1. Percentage utilization of woody plant species by elephants in RINP
Plot
name Name of plant species Utilization/% PART AFFECTED
1
Chaetacme aristata
5
Leaves
1
Ekebergia capensis
5
Stem
1
Ficus sycomoras
2
Stem
1
Lecaniodiscus fraxinifolis
2
Stem
2
Annona senegalensis
2
Stem
3
Lannea fulva
1
Stem
3
Annona senegalensis
50
Stem
3
Phoenix reclinata
30
Stem
3
Phoenix reclinata
30
Leaves
3
Periploca linearifolia
50
Stem
4
Chaetacme aristata
80
Leaves
4
Croton macrostachyus
60
Leaves
4
Salacia madagascarensis
10
Stem
5
Croton sylvaticus
50
Leaves
6
Croton sylvaticus
40
Leaves
6
Chaetacme aristata
30
Leaves
7
Ekebergia capensis
80
Stem
7
Croton sylvaticus
50
Leaves
8
Phoenix reclinata
80
Stem
9
Saba comorensis
30
Stem
10
Croton macrostachyus
10
Leaves
10
Croton macrostachyus
10
Stem
11
Antiaris toxicaria
20
Stem
11
Croton macrostachyus
25
Stem
12
Croton macrostachyus
25
Leaves
13
Chaetacme aristata
20
Leaves
14
Aeschynomene elaphroxylon
30
Stem
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14
Croton macrostachyus
15
Leaves
15
Lecaniodiscus fraxinifolis
40
Stem
15
Croton sylvaticus
50
Leaves
16
Aeschynomene elaphroxylon
60
Stem
17
Aeschynomene elaphroxylon
50
Leaves
17
Saba comorensis
10
Stem
18
Senna spectabilis
40
Leaves
19
Haplocoelum inopleum
10
Leaves
20
Croton sylvaticus
60
Leaves
21
Saba comorensis
50
Stem
21
Landolphia buchananii
10
Stem
22
Aeschynomene elaphroxylon
40
Leaves
23
Croton macrostachyus
80
Leaves
23
Haplocoelum inopleum
10
Leaves
23
Ekebergia capensis
20
Stem
24
Margaritaria discoidea
10
Stem
24
Ekebergia capensis
20
Stem
25
Ficus lutea
50
Stem
25
Phoenix reclinata
45
Stem
25
Phoenix reclinata
45
Leaves
25
Albizia gummifera
20
Leaves
26
Croton macrostachyus
50
Leaves