Biodiversity surveys of Murchison Falls Protected Area.

Abstract

This report summarises the findings of a biodiversity survey of Murchison Falls Protected Area (MFPA - including Murchison Falls National Park, Bugungu and Karuma Wildlife Reserves). The survey shows that MFPA is rich in species and biodiversity with a total known list of 144 mammal species, 556 bird species, 51 reptile species, 28 known amphibian species with an additional 23 to identify (51 species), and 755 plant species. This makes this conservation area relatively biodiverse in the region although not as diverse as some of the parks in further south in the Albertine Rift, such as Queen Elizabeth and Virunga National Parks which also have savannah ecosystems. These surveys have greatly increased the known number of species for the conservation area, doubling the previously known number of reptile species and significantly increasing the number of mammal, bird and plant species from previously published lists. It is likely therefore that with extra survey effort more species could be found given the short duration of our surveys.

While species richness and diversity is high the number of species that are restricted range (Albertine Rift endemic) or globally threatened on the IUCN redlist are not so many and mostly restricted to the mammals, birds and plants. It is potentially possible that the amphibian species that are currently being identified using genetic material may prove to be species of conservation concern or even new species. Of the species we have found for MFPA one mammal is Endangered and four mammals are Vulnerable; four Endangered and seven Vulnerable bird species; two amphibians are Data Deficient, and eight Vulnerable and seven Albertine Rift endemic plant species.

Bugungu Wildlife Reserve was particularly rich for certain taxa, notably birds, amphibians and plants, proving to be richer than Murchison Falls National Park for these taxa. It also contained many of the endemic and threatened plant species. This is likely a result of its varied habitats and its proximity to Budongo Forest but identifies it as being a place of particular conservation value in the landscape.

Full text

Biodiversity Surveys of
Murchison Falls Protected Area
Murchison Falls. A.J.Plumptre/WCS
A. J. Plumptre, S. Ayebare, H. Mugabe, B. Kirunda,
R. Kityo, S. Waswa, B. Matovu, S. Sebuliba, M. Behangana,
R. Sekisambu, P. Mulondo, T. Mudumba,
M. Nsubuga, S. Isoke, S.Prinsloo and G. Nangendo
August 2015

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EXECUTIVE SUMMARY
This report summarises the findings of a biodiversity survey of Murchison Falls Protected Area
(MFPA - including Murchison Falls National Park, Bugungu and Karuma Wildlife Reserves). The
survey shows that MFPA is rich in species and biodiversity with a total known list of 144 mammal
species, 556 bird species, 51 reptile species, 28 known amphibian species with an additional 23 to
identify (51 species), and 755 plant species. This makes this conservation area relatively biodiverse in
the region although not as diverse as some of the parks further south in the Albertine Rift, such as
Queen Elizabeth and Virunga National Parks which also have savannah ecosystems. These surveys
have greatly increased the known number of species for the conservation area, doubling the
previously known number of reptile species and significantly increasing the number of mammal, bird
and plant species from previously published lists. It is likely therefore that with extra survey effort
more species could be found given the short duration and limited sampling of our surveys.
While species richness and diversity is high, the number of species that are restricted range (Albertine
Rift endemic) or globally threatened on the IUCN global redlist are not so many and mostly restricted
to the mammals, birds and plants. It is potentially possible that the amphibian species that are
currently being identified using genetic material may prove to be species of conservation concern or
even new species. Of the species we have found for MFPA there are two Endangered and four
Vulnerable mammal species; four Endangered and seven Vulnerable bird species; two amphibians are
Data Deficient, and eight Vulnerable and seven Albertine Rift endemic plant species.
Bugungu Wildlife Reserve was particularly rich for certain taxa, notably birds, amphibians and plants,
proving to be richer than Murchison Falls National Park for these taxa. It also contained many of the
endemic and threatened plant species. This is likely a result of its varied habitats and its proximity to
Budongo Forest, but identifies it as being a place of particular conservation value in the landscape.
ACKNOWLEDGEMENTS
We would like to thank many people who helped WCS implement these surveys. The Conservation
Area Manager of Murchison Falls Protected Area, Mr. Tom Okello was very helpful in organizing
logistics for the field teams and providing advice on where teams could access safely. We are grateful
to the field assistants who collected the data for these surveys, particularly: Moses Gonya, Nabert
Mutungire, Julius Kyamanywa, Conslate Alezuyo, Wilber Lukwago and Akoth Sisiria and to the
UWA rangers who also took part in the data collection. We also thank Richard Musumererwa Amooti
for ably cooking for all the field teams, and Sam Mulondo, Bosco Kirama and Wilson Muhumuza for
driving the survey vehicles. Robert Kityo at the Makerere University Museum identified the small
mammals for this survey and Mathias Behangana identified the reptiles and amphibians.
We want to thank The Norwegian Government for the financing of this work through NEMA and
UWA and Tullow Oil for their financial support for the surveys in Bugungu WR and in the human
modified areas. We are grateful to Uganda Wildlife Authority for permission to carry out the surveys
in the Murchison Falls Protected Area.

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TABLE OF CONTENTS
EXECUTIVE SUMMARY .................................................................................................................................. 2
ACKNOWLEDGEMENTS ................................................................................................................................. 2
TABLE OF CONTENTS ..................................................................................................................................... 3
INTRODUCTION ................................................................................................................................................ 4
METHODS ............................................................................................................................................................ 6
Survey Design .................................................................................................................................................... 6
Large mammal surveys ...................................................................................................................................... 6
Small mammal surveys ....................................................................................................................................... 6
Amphibian and reptile surveys ........................................................................................................................... 7
Bird Surveys ....................................................................................................................................................... 7
Plant surveys ...................................................................................................................................................... 8
RESULTS .............................................................................................................................................................. 9
Large mammals.................................................................................................................................................. 9
Small mammals .................................................................................................................................................. 9
Birds ................................................................................................................................................................. 13
Reptiles ............................................................................................................................................................ 16
Amphibians ...................................................................................................................................................... 17
Plant species .................................................................................................................................................... 19
CONSERVATION IMPLICATIONS............................................................................................................... 23
Biodiversity and species of Conservation Concern in MFPA .......................................................................... 23
Conservation of MFPA .................................................................................................................................... 23
REFERENCES ................................................................................................................................................... 25

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INTRODUCTION
The Murchison Falls Protected Area (MFPA) includes the Murchison Falls National Park (MFNP -
3,898 km2), Karuma Wildlife Reserve (Karuma WR - 678 km2) and Bugungu Wildlife Reserve
(Bugungu WR - 474 km2). The two wildlife reserves are contiguous with the national park and
together form a conservation unit that encompasses 5,045 km2 of natural habitat, Uganda’s largest
landscape of protected areas (figure 1).
Figure 1. Map of the Murchison Falls Protected Area with the main vegetation types mapped.
When Samuel Baker discovered the falls in 1866 the banks of the Nile River which flows through the
park were heavily populated. Outbreaks of Rinderpest and Sleeping Sickness in the late 1800s led to
the out-migration of people and a government resettlement scheme from 1912 (Olupot et al. 2010).
Murchison Falls National Park (MFNP) was established as the Bunyoro Game Reserve in 1910 and
then gazetted as a national park in 1952. Karuma and Bugungu Controlled Hunting Areas were
established in 1962 to allow sport hunting of elephants and other species and these became Game
Reserves later in the 1960s and finally Wildlife Reserves in 1996 after the formation of the Uganda
Wildlife Authority (UWA) from the merger of the Uganda National Parks and Game Department
(Olupot et al. 2010). Following its creation as a national park, Murchison Falls became a significant
tourism attraction in East Africa with more than 60,000 visitors coming to the park in the late 1960s
and early 1970s.
MFPA was probably the hardest hit of any of the protected areas in the civil unrest of the 1970s and
1980s. Elephant numbers declined from 14,000 in the late 1960s to 250 in 1983/84. It was only
following the establishment of the current government in Uganda and the stabilization of the rule of
law that elephant and other large mammal numbers started to increase again. An aerial survey in 2014
by the Wildlife Conservation Society (WCS) and UWA showed Murchison’s elephants now number
about 1,300. The loss of so many elephants has led to the expansion of woodland and forest within
MFPA (figure 1).

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Research in the MFNP focused on large mammal ecology, particularly the ‘problem of elephants’
which occurred as elephants lost habitat outside the MFPA resulting in migration and a high density
of elephants within the park. This led to major habitat alteration with the conversion of wooded areas
to grassland. Culling of elephants over the years to reduce human-wildlife conflict allowed detailed
research to be made on the reproduction and population dynamics of this species (Laws, Parker and
Johnstone, 1975). No formal surveys of the biodiversity of the park were made however.
Published records for MFNP (Wilson 1995) showed the park to contain in the region of 76 mammal
species, an observation that was probably an under-estimate given that surveys for small mammal
species (bats, rodents and shrews) have only been conducted for some sections of the park, and 450
bird species (19 unconfirmed at the time). Biodiversity surveys were made of Bugungu Wildlife
Reserve (Allan 1997) which focused on large and small mammals, birds, reptiles, amphibians and
invertebrates. This survey recorded 227 birds for Bugungu WR (33 not seen in MFNP), 26 large
mammals, fourteen species of rodent, five species of shrew and five species of bat (total of 50
mammal species). Of the ungulates a key finding was the red-flanked duiker (Cephalophus rufiliatus)
in Bugungu WR which may not occur elsewhere in Uganda. Reptiles numbered 26 and amphibians 15
species with some remaining to be identified at the time of the report. 47 butterfly and 9 dragonfly
species were also identified.
However, there has never been a comprehensive biodiversity assessment for MFPA as a whole. There
have been some lists generated from surveys made for EIA’s and some basic baseline surveys for the
oil companies but nothing that has attempted to cover the whole conservation area. In 2014 WCS was
requested by UWA to help them undertake a survey of the biodiversity of MFPA to provide a baseline
for monitoring of future changes in the biodiversity of the landscape. Oil and gas exploration in
MFPA has shown that there are significant reserves under the park and wildlife reserves and these are
very likely to be exploited in the future. The Norwegian Government funded the Government of
Uganda to support preparations for oil and gas production, part of which included support to baseline
surveys. This report summarises the results of the baseline biodiversity surveys for MFPA. At the
same time Tullow Oil was funding WCS to undertake baseline biodiversity surveys of Exploration
Area 2 which included part of Bugungu Wildlife Reserve as well as the community land to the north
and south west of this area. We have therefore included the results for Bugungu Wildlife Reserve
surveys funded by Tullow Oil in this report.
The biodiversity surveys focused on terrestrial vertebrates and plants; mammals, birds, reptiles,
amphibians and higher plants. While it would have been good to include other taxa the funds available
for these surveys were limited and as a result we focused on these taxa which can be identified
relatively easily because of good keys. While one objective was to compile a list of species for the
park, the key objective was to map the diversity spatially across the park to highlight areas of species
richness or that are important for threatened species.

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METHODS
Survey Design
The software DISTANCE 6.0 was used to design a survey using a stratified-random sampling method
of 3 km transects across the MFPA (Figure 2). Spacing of the transects was determined by the
available budgets with closer spacing in Bugungu Wildlife Reserve and in human modified habitat in
Buliisa District because of Tullow Oil funding for that site. Bird and plant teams aimed to reach as
many transects as possible, given difficulties in accessing parts of MFPA, while reptile/amphibian and
small mammal teams targeted fewer transects but ensured that the diversity of habitats were sampled.
Six additional transects were added to sample rarer habitats in the Bugungu area and four transects
that were initially planned had to be re-located because of difficulty of access.
Figure 2. The design of transects surveyed across MFPA showing both transects selected in
DISTANCE design and additional ones added to sample specific habitat.
Large mammal surveys
UWA undertakes regular aerial surveys of large mammals in MFPA with the last survey made with
WCS in June 2014. There have also been records made over the years of sightings of large mammal
species. As a result the large mammal fauna is relatively well known and didn’t need to be
resurveyed. However, WCS and UWA both had camera traps available and WCS placed 55 camera
traps across MFPA to look at relative capture rates of different species with a particular interest in
species that are rarely observed.
Small mammal surveys
Shrews/Rodents
Capturing of different mammal species requires different trapping techniques and wide variety of
traps for trapping small terrestrial mammals are available. For these surveys however Sherman traps
were used to capture rodents and shrews, using a bait that comprised of a mixture of peanut butter,
maize flour, margarine and bananas- a usually effective bait for trapping a wide spectrum of small

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terrestrial mammals. The trapping protocol used traps laid along line transects that maximized the
habitat variation in each survey area. To enhance the chances of capturing animals, traps were
specifically placed at locations with feeding signs, runways and against or beneath logs and areas with
a good amount of low vegetation cover.
All trap locations were marked with a GPS and also marked with flagging tape so that traps could be
easily relocated. For the rodents and shrews, a uniform trap effort was used for all survey areas.
Bats
Bats were sampled using mist nets, searching for roosts, and using an acoustic bat detector that can
record micro-chiropteran activity. Sampling of bats was constrained by the location of sampling sites
and the safety of the field team at night and as a result not every site sampled for shrews and rodents
was sampled for bats also.
Amphibian and reptile surveys
In each of the main habitat types of the MFPA, reptiles and amphibians were surveyed using Visual
Encounter Surveys (VES) and Pitfall Traps. These methods were used to document the presence of
amphibians and reptiles and sample individuals for each species.
Visual Encounter Surveys
Visual Encounter Surveys (VES) are a well known and robust method for survey hepterofauna. VES
is similar to the Timed Constrained Count (TCC) method described by Heyer et al., (1994). Visual
encounter surveys are used to document presence of amphibians and reptiles and are effective in most
habitats and for most species that tend to breed in lentic habitats. They generate encounter rates of
species in their habitats in a unit hour.
The method comprises moving through a habitat, turning logs or stones, inspecting retreats and
watching out for and recording surface-active species. The data gathered using this procedure
provides information on species richness of the habitat. For amphibian fauna, the best results are
achieved when the surveys take place in the evenings between 1900 and 2100 hours as this is when
most amphibians are active. For reptiles, there is no particular time for sampling all reptiles because
the different groups are active at different times of the day and night. For example, whereas most
tortoises, skinks, agamids and some geckoes are active during the warm parts of days, other species of
geckoes and snakes are nocturnal. Surveying reptiles therefore was more habitat based than temporal.
Pitfall trapping with drift fence
Pitfall traps were set up with a drift fence in selected habitats to sample any surface dwelling
herpetofauna. The use of drift fences with bucket pitfall traps has been the commonest technique for
studies of individual species or herpetofaunal communities and has been used with success for
amphibians (Mitchell et al., 1993; Heyer et al., 1994, Handley and Varn, 1994; Msuya, 2001). The
results of studies employing drift fences with pitfall traps provide valuable insights into population
and community ecology, and behavioural patterns of secretive and difficult to study species (Dodd,
1991). Each drift fence comprised of 10, 20-litre plastic buckets placed at an interval of 10 m,
covering a total length of 100 m. The buckets were placed in holes dug in the substrate such that their
rim was level with the ground. A 100-meter long and 0.5 m high drift fence of black polythene
supported vertically by wooden laths was set in an alternating manner with the buckets in the line to
permit detection of directional movement of herpetofauna. The pitfall traps were inspected twice a
day.
Bird Surveys
Birds were sampled using point counts at 250 m along each transect visited and the habitat of each
point noted. Additional points were placed in rare habitats in the vicinity of the sample site. The latter

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would include feeding sites for migratory waders, small wetlands etc. Point counts consisted of the
two ornithologists arriving at a site and waiting a minute for the birds to settle down if they had been
disturbed. They would then make a five minute point count noting all birds seen or heard from the
point and the distance from the observer to the bird in the following distance classes: 0-10m, 10-20m,
20-50m, 50-100m, 100-200m, 200+m.
The ornithologists working with WCS can identify all the bird calls from birds that were likely to be
found in MFPA and also aimed to compile a total species list for each transect in addition to the
quantitative data obtained from the point counts.
Plant surveys
Plant plots were measured at 250 metres along each transect visited. Standard nested circular plots
have been used at all sites in the AR. Small herbs are recorded in a circle of 2m radius; trees from 2.5-
10cmDBH, lianas (>1cm diameter) and shrubs are recorded in 10m radius plots; and trees >
10cmDBH are recorded in 20m radius plots. A representative sample was taken of every species
identified in the field so that checks could be made on species identifications later. A GPS reading
was made for every plot and habitat, slope and canopy cover measurements were made using a
standard form.
A total list of species at each transect was compiled by collecting fertile plant specimens (flower/fruit)
where possible and non-fertile otherwise of all plant species detected. The specimens were pressed
and dried using portable plant driers that WCS has developed for field surveys. Specimens were
identified at Makerere University Herbarium. In addition, for each specimen a GPS point and field
notes on habitat and characteristics were recorded.

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RESULTS
Large mammals
A total of 48,824 photographs were taken by camera traps placed in the MFPA during these surveys.
From these 38 large or medium sized mammal species were observed (Table 1). Few cameras were
placed in Karuma WR so no data are given for here. While these species have all been recorded
previously for the MFNP area it is encouraging that there are still species such as the Giant Pangolin,
Ratel (Honey badger) and the Bunyoro rabbit which is only found from here to north eastern
Democratic Republic of Congo.
Table 1. List of medium sized mammals observed in the MFPA from camera trap photos
Order Species MFNP Bugungu
Primate
Black and white colobus
1
Chimpanzee
1
Olive Baboon
1
1
Patas monkey
1
Vervet
1
Carnivora
African Civet
1
Large spotted genet
1
1
Rusty-spotted genet
1
Servaline Genet
1
1
Banded mongoose
1
Marsh Mongoose
1
1
Slender mongoose
1
White tailed mongoose
1
Serval cat
1
Lion
1
Leopard
1
Side striped Jackal
1
Spotted hyena
1
1
Ratel
1
1
Pholidota
Giant Pangolin
1
1
Artiodactyla
Buffalo
1
1
Bush Duiker
1
1
Bush pig
1
1
Bushbuck
1
1
Rothschild Giraffe
1
Hippopotamus
1
1
Jackson's Hartebeest
1
Oribi
1
Reedbuck
1
Uganda kob
1
1
Warthog
1
1
Waterbuck
1
1
Proboscidae
Elephant
1
1
Lagomorpha
Bunyoro Rabbit
1
Rodentia
Cane rat
1
Crested porcupine
1
1
Tubulidentata
Aardvark
1
1
Small mammals
A total of twenty one sites were visited where a camp was established and then surveys made in the
vicinity of the camp. Of the 317 small mammals trapped/encountered, 60 small mammal species were

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recorded from the three protected areas in the MFPA. These included 27 bat species, 15 shrew
species, one rabbit (Poelagus marjorita), one galago (Galagoides thomasi) and 16 rodent species.
Bats
Table 2 gives the 27 bat species encountered during these surveys and the number observed or trapped
in each of the three protected areas within MFPA. It can be seen from these results that MFNP is
particularly rich in bat species compared with the Wildlife Reserves. One species of bat found on
these surveys, Chaerophon russatus, was the first record for Uganda for this species.
Table 2. The number of each species of bat observed/trapped in the three protected areas in MFPA.
Family Row Labels Bugungu Karuma MFNP
Fruit Bats
Pteropodidae
Epomophorus labiatus 8 1 12
Epomops franqueti
3
1
Hypsignathus monstrosus
1
Micropteropus pusillus
4
1
Rousettus angolensis
1
Insectivorous bats
Nycteridae Nycteris macrotis
1
Nycteris thebaica
20
Hipposideridae
Hipposideros ruber
9
Megadermatidae
Lavia frons
6
Embalonuridae Taphozous mauritianus
4 1
Vespertilionidae
Glauconycteris argentata
4
Glauconycteris humeralis
1
Glauconycteris variegata
1
Mimetillus moloneyi
1
Pipistrellus capensis
1
4
Pipistrellus guineensis
1
Pipistrellus nanulus
1
Pipistrellus nanus
2
Pipistrellus rueppelli
1
Pipistrellus somalicus
2
Scotoecus albofuscus
2
Scotoecus hirundo
4
Scotophilus dinganii
4
Molosidae
Chaerephon ansorgei
2
Chaerephon pumilus
1
Chaerophon russatus*
1
Mops condylurus
3
Number of species 3 6 24
*This is the first record of this species for Uganda

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Rodents and Shrews
The 16 rodent species are listed in table 3 together with the 15 shrew species. The number of rodent
species was more similar between protected areas and for shrews Bugungu and MFNP were similar in
number but Karuma had few species. One rodent species, Thamnomys venustus, is Vulnerable on the
IUCN global redlist and is also an Albertine Rift endemic species. It was found in Karuma WR.
Table 3. The number of shrew and rodent species observed in the three protected areas in MFPA.
Shrews
Bugungu
Karuma
MFNP
Crocidura denti
2
Crocidura dolichura
1
Crocidura fuscomurina
1
Crocidura gracilipes
1
2
Crocidura hildegardeae
5
Crocidura hirta
2
Crocidura jacksoni
1
Crocidura luna
2
1
Crocidura nanilla
1
Crocidura nigrofusca
8
Crocidura olivieri
4
4
Crocidura parvipes
3
Crocidura roosevelti
2
Crocidura sp
6
4
11
Crocidura turba
3
Total Shrew species
8
2
10
Rodentia
Bugungu
Karuma
MFNP
Aethomys hindei
3
3
9
Aethomys kaiseri
2
4
Dendromus melanotis
1
Dendromus mystacalis
1
1
Grammomys dolichurus
2
1
Graphiurus murinus
2
Lemniscomys barbarus
1
1
1
Lemniscomys macculus
1
Lemniscomys striatus
4
6
11
Lophuromys aquilus
6
1
Lophuromys sikapusi
1
3
2
Mastomys natalensis
19
24
Mus mahomet
5
1
10
Mus musculoides
5
2
6
Mus triton
2
Thamnomys venustus
1
Total Rodent species
11
10
12
Sampling intensity differed between protected areas and a comparison between sites is best made with
rarefaction curves (Figure 3). This shows that MFNP was richer in small mammal species compared
with the two Wildlife Reserves which were similar in small mammal species richness. Mapping the

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relative distribution of small mammal species richness did not show any clear pattern across MFPA
(figure 4).
Figure 3. Rarefaction curves for the three protected areas showing the greater species richness of
MFNP.
Figure 4. Map of relative species richness of small mammals across MFPA.

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Species diversity was calculated for small mammals in each protected area (table 4) and shows the
higher diversity in MFNP whether using the Shannon-Wiener or Alpha index.
Table 4. Shannon Wiener diversity and evenness and the Alpha diversity index calculated for the
three protected areas.
Index
Bugungu Wildlife
Reserve
Karuma Wildlife
Reserve
Murchison Falls
Shannon H' Log Base
10.
1.134
1.177
1.469
Shannon J' (evenness)
0.833
0.938
0.874
Alpha diversity index
9.524
10.664
21.992
Birds
A total of 340 bird species were recorded in the MFPA with 133 species recorded in Karuma WR, 232
in Bugungu WR, 220 in Murchison Falls National Park and 162 in the human modified landscape
north and south of Bugungu. The lower numbers of species for Karuma WR were mainly due to a
lower sampling effort there compared with the other two protected areas. A total of 556 bird species
are known from MFPA, adding 106 species to the list of Wilson (1995), mostly forest species from
the Wildlife Reserves. Rarefaction curves, which plot the number of species observed against number
of birds seen, show that the species richness is likely to be slightly higher in Bugungu WR than
MFNP. The human modified landscape had many fewer species and the rarefaction curves had almost
leveled off at 162 (figure 5).
Figure 5. Rarefaction curves for the three sites plotted against the number of birds observed. Data
were from point counts.
The distribution of species richness indicates that the number of species seen at each of the transects
varies from 19 to 72 (figure 6). Some areas of Karuma Wildlife Reserve and northern MFNP had few
species despite several point counts at each site. The richest areas were in central MFNP and also at
the edge of Bugungu and MFNP where transects were also in the community land (human modified

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habitat). Species that are found in human modified habitat tend to be common species but will
increase species richness values of these sites.
Figure 6. Relative bird species richness from point counts along each transect surveyed in the three
protected areas.
Figure 7. Locations of sightings of threatened bird species.

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No bird species that are endemic to the Albertine Rift were observed in the MFPA but four threatened
species were; African White-backed Vulture (Gyps africanus) and Ruppell’s Vulture (Gyps ruppelli)
which are Endangered and were only recorded in MFNP (at 5 point counts each) and not in the
wildlife reserves, Martial Eagle which is Vulnerable was recorded at five point counts; two in MFNP
and two in Bugungu WR and one in Karuma WR; and Grey-crowned Crane (Balearica pavonina)
which is endangered and was recorded at seven point counts. Figure 7 plots the locations of these
sightings and shows that only one threatened species was seen at most transects but two threatened
species were seen on two transects in the heart of the park.
Diversity measurements using the Shannon Wiener Diversity measurement and the Log Series Alpha
diversity measurement show that Bugungu tends to be the most diverse site (table 5). The Shannon-
Wiener index is presented here because it is often used in studies and so allows comparisons to be
made with other studies. However, it is known to overweight rare species and that the alpha diversity
index is a better measurement of diversity (Krebs 1989).
Table 5. Shannon Wiener diversity and evenness and the Alpha diversity index calculated for the
three protected areas.
Index
Bugungu Wildlife
Reserve
Murchison Falls
Human
modified
habitat
Karuma Wildlife
Reserve
Shannon H' Log Base
10.
1.85
1.77
1.74
1.74
Shannon J' (evenness)
0.78
0.76
0.79
0.82
Alpha diversity index
48.07
41.53
29.35
39.44
A cluster analysis using the Bray Curtis similarity measure showed that the bird community in human
modified habitat in Buliisa District was most similar to the bird community in Bugungu Wildlife
Reserve (Figure 8). Karuma Wildlife Reserve had a very different bird community, mainly because of
the dense woodland that is found there, with less than 25% overlap in species composition with the
other three sites. These results show that each protected area is conserving different communities of
birds and despite their close juxtaposition the two wildlife reserves are important in their own right for
the diversity of MFPA.
Figure 8. The percentage similarity in bird fauna between the three protected areas. Despite similar
numbers of species there are big differences in species composition.

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Reptiles
A total of 29 reptile species were recorded from these surveys. Eighteen species were found in
Bugungu WR, six species in Karuma WR and 25 species in MFNP. None of these were endemic to
the Albertine Rift or threatened species. This brings the list of reptiles for the MFPA to 51 compiling
prior lists from surveys of this region and these surveys, including 44 species in MFNP, 25 in
Bugungu WR and 18 in Karuma WR. No species endemic to the Albertine Rift or threatened species
are known form MFPA.
Reptiles and amphibians are collected opportunistically and only one or two specimens are collected
for identification purposes at any site. It is therefore not possible to plot rarefaction curves using
number of individuals encountered at a site on the X-axis. Instead we plotted number of sites with
number of species at a site (figure 9).
Figure 9. Rarefaction curve for the three protected areas. Karuma only has one point as few sites were
sampled.
Figure 10. Relative richness of reptile species across MFPA.

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Reptile species were more numerous in the western side of MFNP and in sites in Bugungu WR when
compared with the eastern side of MFNP (figure 10).
The diversity of reptile species as measured by Shannon-Wiener or the alpha index differ by site
(table 6). With the alpha measure the two wildlife reserves are more species rich while with the
Shannon-Wiener index MFNP is slightly richer. Given the close rarefaction curves in figure 9 and the
low sampling effort in Karuma WR it is likely that with more effort better measures of diversity
would be obtained.
Table 6. Shannon Wiener diversity and evenness and the Alpha diversity index calculated for the
three protected areas.
Index
Bugungu Wildlife
Reserve
Murchison Falls
Karuma Wildlife
Reserve
Shannon H' Log Base
10.
1.169
1.239
0.759
Shannon J' (evenness)
0.931
0.886
0.976
Alpha diversity index
17.926
12.481
19.941
Amphibians
A total of 45 amphibian species were found in the MFPA. Amphibians are difficult to identify to
species on morphology alone and samples have been sent to the Trento Science Museum for genetic
analysis to confirm preliminary species identifications. Some species however, could only be
identified to genus level. A total of 36 species were found in MFNP, 30 in Bugungu WR and 13 in
Karuma WR but the later was only sampled at two locations. No Albertine Rift endemic species nor
threatened species were found. Of these species 23 still need to be identified to species level from
Genus level.
Figure 11. Rarefaction curves for the three sites plotted for the number of sites with species.

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The rarefaction curves show that Bugungu WR looks to be more species rich than MFNP or Karuma
WR, although the sampling of only two sites in Karuma limits what we can determine for this site.
No clear pattern in richness occurs across the MFPA (figure 12) but higher richness seems to be in the
wooded and forested areas.
Figure 12. Relative richness of amphibian species across MFPA.
Table 7. Shannon Wiener diversity and evenness and the Alpha diversity index calculated for the
three protected areas.
Index
Bugungu Wildlife
Reserve
Murchison Falls
Karuma Wildlife
Reserve
Shannon H' Log Base
10.
1.419
1.43
0.954
Shannon J' (evenness)
0.912
0.968
1
Alpha diversity index
14.838
37.416
7.162
Both Shannon-Wiener and alpha diversity indices rank MFNP as being richer than the wildlife
reserves, probably because of the fewer sampling sites. The rarefaction curves give a better
comparison of sites (figure 11).

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Plant species
A total of 755 plant species were identified from 455 plots surveyed across all sites with 539 species
in Bugungu WR, 421 species for MFNP, 303 species in the human modified habitat and 275 species
in Karuma WR. Bugungu WR is significantly richer in plant species as shown in the rarefaction
curves while the other sites were all similar in species richness (figure 13).
Eight threatened plant species were recorded (all globally vulnerable) with five in Bugungu WR, three
in Karuma WR, two in MFNP and one in the Human modified habitat in Buliisa (Table 8). Bugungu
not only has more species of threatened plant but also had more plots with threatened species. Seven
species endemic to the Albertine Rift were also found with four in Bugungu WR and only one at each
of the other sites. Bugungu also had more plots with endemic species found in them (Table 9).
Figure 13. Rarefaction curves for plant species. Data from plots.
Table 8. The number of plots in which threatened species of plant (VU) were found in each of the
three protected areas and human modified habitat.
Species
Bugungu WR
Human modified
habitat
Karuma WR
Murchison Falls
National Park
Afzelia africana
3
Albizia ferruginea
1
Entandrophragma
cylindricum
2
Khaya anthotheca
4
1
Khaya grandifoliola
3
1
Lovoa trichilioides
2
Prunus africana
1
Psilotrichum axilliflorum
4
1
Number of plots with
threatened species
14
1
4
4

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Table 9. The number of plots in which species of plant endemic to the Albertine Rift were found in
each of the three protected areas and human modified habitat.
Species
Bugungu WR
Human modified
habitat
Karuma WR
Murchison Falls
National Park
Coccinia mildbraedii
2
Entada phaneroneura
12
Isoglossa laxiflora
3
Isoglossa vulcanicola
2
Rytigynia bugoyensis
1
Thunbergia
mildbraediana
2
Tinospora orophila
1
Number of plots with
threatened species
18
1
2
2
Figure 14. Plant species richness from plots along each transect.
Figure 14 shows the species richness of plants for each transect surveyed across the MFPA. It is clear
the highest species richness is found on transects close to the Budongo Forest and in colonising forest
(compare figure 14 with Figure 1). Grassland areas in MFNP have a very low species richness which
is why the park ranks so much lower than Bugungu WR in terms of total species richness because
grassland cover is much greater in the park.
Figures 15 and 16 plot the number of threatened plants and plants endemic to the Albertine Rift
respectively. These figures show how important Bugungu WR is for endemic plant species which
were found in forest and along the escarpment above Lake Albert. Threatened plant species appear to

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be closely associated with forested or dense bushland areas and hence were more common in the
forests in Bugungu WR.
Figure 15. Richness of plant species that are VU under IUCN global redlisting
Figure 16. Number of Albertine Rift endemic plant species found on each transect

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Diversity measurements using the Shannon Wiener Diversity measurement and the Log Series Alpha
diversity measurement show that Bugungu WR tends to be the most diverse site although the Shannon
Wiener evenness index indicates that Karuma WR is more even in its composition of species (table
10). The alpha diversity index ranks the human modified habitat as far less diverse than the Shannon
Wiener index. The Shannon-Wiener index is presented here because it is often used in studies and so
allows comparisons to be made with other studies. However, it is known to overweight species that
are rarely encountered and that the alpha diversity index is a better measurement of diversity (Krebs
1989).
Table 10. Shannon Wiener diversity and evenness and the Alpha diversity index calculated for the
three protected areas.
Index
Bugungu Wildlife
Reserve
Murchison Falls
Human
modified
habitat
Karuma Wildlife
Reserve
Shannon H' Log Base
10.
1.85
1.77
1.74
1.74
Shannon J' (evenness)
0.78
0.76
0.79
0.82
Alpha diversity index
48.07
41.53
29.35
39.44
A comparison of the similarity between communities using a Bray-Curtis clustering showed that
Bugungu WR and MFNP were the most similar in species composition with 47% overlap but that
Karuma WR was the most dissimilar with less than 30% overlap with any of the other sites (figure
17).
Figure 17. The percentage similarity in plant community between the three protected areas and
human modified habitat.

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CONSERVATION IMPLICATIONS
Biodiversity and species of Conservation Concern in MFPA
Combining the species described form the surveys above with previously published species lists for
the park brings the numbers of species to many more than previously published (Table 11). Wilson
(1995) published 76 mammal species for Murchison Falls National Park which has increased to 144
for MFPA with these surveys as well as other surveys that have been made in the park and wildlife
reserves. This increase is mostly due to the intensive surveys of small mammals, particularly the
shrews and bats. Similarly plant species have increased from 450 (J. Kalema in grey literature) to 755
with some specimens remaining to be identified. Bird numbers increased form 450 (Wilson 1995) to
555 with this survey as well as the Frontier surveys of Bugungu WR (Allen 1997). Plant species of
conservation concern are listed in Tables 8 and 9 and threatened vertebrates are listed in table 12.
Table 11. The number of species for each taxon and number of restricted range species (Albertine
Rift Endemic) and threatened species known from MFPA.
Taxon
Number of species
Albertine Endemics
Number threatened
Mammals
144
1 EN; 4 VU
Birds
556
4 EN; 7VU
Reptiles
51
Amphibians
51
2 DD
Plants
755
7
8VU
Table 12. Threatened vertebrates from MFPA
Mammals
Birds
Amphibians
Chimpanzee (EN)
African White-backed Vulture (EN)
Bufo vitattus (DD)
Rothschild giraffe (EN)
Ruppell's Vulture (EN)
Ptychadena chrystyi (DD)
Lion (VU)
Hooded Vulture (EN)
Hippopotamus (VU)
Egyptian Vulture (EN)
Elephant (VU)
Shoebill (VU)
Thamnomys venustus (VU)
Secretary Bird (VU)
Lappet-faced Vulture (VU)
White-headed Vulture (VU)
Martial Eagle(VU)
Lesser Kestrel (VU)
Black-crowned Crane (VU)
Conservation of MFPA
The results of the surveys greatly increased the conservation value of this landscape, increasing
species richness and identifying more species of global conservation concern. It also highlighted the
biological value of Bugungu Wildlife Reserve which in the case of birds, amphibians and plants was
richer in species than Murchison Falls National Park (see rarefaction curves). It was also the site
where many of the endemic and threatened plants occurred. This is probably because Bugungu is very
diverse in habitats and also includes Tropical High Forest with its overlap with the Budongo Forest
Reserve.

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Combining the species richness maps for all five taxa across MFPA is possible by calculating the
ranking of each survey site (transect) in relation to other sites in terms of the number of species
encountered for each taxon separately. The mean ranking is then calculated across the five taxa to
provide an average value of species richness across the taxa (figure 18). This shows the importance of
Bugungu WR graphically as well as the central areas of MFNP. There are rumours currently that
politicians are lobbying for the degazettment of parts of Bugungu to provide land for people in the
Buliisa District. In-migration of people looking for work in the Oil Industry as well as the collapse of
the fisheries on Lake Albert is leading to limited livelihood options and the desire for land for
cultivation. We would argue that these results show the biological importance of Bugungu WR for
conservation.
Interestingly the species richness of sites outside the protected areas was also relatively high (mainly
bird and plant species). Some of these sites were remaining wetlands and patches of natural habitat
surrounded by farmland, but in other cases species richness could be high for bird species but they
tended to be species that were very common and not of conservation value.
Figure 18. Average ranking across taxa, highlighting the areas of high overall biodiversity.
The development of Oil production in this region is of concern for the biodiversity that we report
here. While we believe the oil companies and Government are making the correct approaches,
working to minimize the impacts of the oil industry on this landscape these results make it even more
imperative that great care is taken to ensure that none of the biodiversity is lost as a result of these
activities. Careful planning of pipeline routes and roads should be made to avoid passing through the
protected areas and where possible they should pass along the border of the protected areas and
should be fenced. This would also reduce crop raiding conflict between the local people and Uganda
Wildlife Authority and thereby encourage a win-win situation. However exact routing and fence
locations and the manner of construction must still allow for migration of species between the MFPA
and other important biodiversity sites, particularly to Budongo Forest, along the escarpment edges and
forest corridors.

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