The Biodiversity of the Albertine Rift

Abstract

The Albertine Rift is one of the most important regions for conservation in Africa. It contains more vertebrate species than any other region on the continent and contains more endemic species of vertebrate than any other region on mainland Africa. This paper compiles all currently known species distribution information for plants, endemic butterfly species and four vertebrate taxa from the Albertine Rift. The literature on fish species richness and endemism is also reviewed to assess the importance of the larger lakes in the Rift for conservation. We use data from 38 protected and unprotected areas to prioritise sites within the Albertine Rift for conservation based upon their numbers of endemic and globally threatened species. Virunga and Kahuzi Biega National Parks and Itombwe Massif in Democratic Republic of Congo, Bwindi Impenetrable and Kibale National Parks in Uganda, and Nyungwe National Park in Rwanda rank highest in terms of numbers of both endemic and globally threatened species. Six conservation landscapes are described that include most of these sites and it is argued that a focus on these landscapes may be a more holistic method to ensure the safety of the priority areas of the Albertine Rift.

Full text

The biodiversity of the Albertine Rift
Andrew J. Plumptre
a,
*, Tim R.B. Davenport
b
, Mathias Behangana
c
, Robert Kityo
c
,
Gerald Eilu
c
, Paul Ssegawa
c
, Corneille Ewango
a
, Danny Meirte
d
, Charles Kahindo
c
,
Marc Herremans
d,k
, Julian Kerbis Peterhans
e,f
, John D. Pilgrim
g,l
, Malcolm Wilson
h
,
Marc Languy
i
, David Moyer
j
a
Wildlife Conservation Society, P.O. Box 7487, Kampala, Uganda
b
Wildlife Conservation Society, P.O. Box 1475, Mbeya, Tanzania
c
Makerere University, P.O. Box 7062, Kampala, Uganda
d
Royal Museum for Central Africa at Tervuren, Leuvensesteenweg 11, 3080 Tervuren, Belgium
e
Roosevelt University, University College, 430 S Michigan Avenue, Chicago, IL 60605, USA
f
Field Museum of Natural History, Chicago, 1400 S. Lake Shore Drive, Chicago, IL 60605-2496, USA
g
Center for Applied Biodiversity Science, Conservation International, 1919 M Street NW, Suite 600, Washington, DC 20036, USA
h
P.O. Box 178, Groblersdal 0470, Mpumalanga, South Africa
i
WWF Eastern Africa Regional Programme Office (EARPO), P.O. Box 62440, 00200 Nairobi, Kenya
j
Wildlife Conservation Society, P.O. Box 936, Iringa, Tanzania
k
Natuurpunt.Studie, Coxiestraat 11, 2 800 Mechelen, Belgium
l
BirdLife International in Indochina, 4/209 Doi Can, Ba Dinh, Hanoi, Vietnam
ARTICLE INFO
Article history:
Received 16 June 2005
Received in revised form
17 July 2006
Accepted 8 August 2006
Available online 14 November 2006
Keywords:
Albertine Rift
Priority setting
Mammals
Birds
Reptiles
Amphibians
Plants
Biodiversity
ABSTRACT
The Albertine Rift is one of the most important regions for conservation in Africa. It con-
tains more vertebrate species than any other region on the continent and contains more
endemic species of vertebrate than any other region on mainland Africa. This paper com-
piles all currently known species distribution information for plants, endemic butterfly
species and four vertebrate taxa from the Albertine Rift. The literature on fish species rich-
ness and endemism is also reviewed to assess the importance of the larger lakes in the Rift
for conservation. We use data from 38 protected and unprotected areas to prioritise sites
within the Albertine Rift for conservation based upon their numbers of endemic and glob-
ally threatened species. Virunga and Kahuzi Biega National Parks and Itombwe Massif in
Democratic Republic of Congo, Bwindi Impenetrable and Kibale National Parks in Uganda,
and Nyungwe National Park in Rwanda rank highest in terms of numbers of both endemic
and globally threatened species. Six conservation landscapes are described that include
most of these sites and it is argued that a focus on these landscapes may be a more holistic
method to ensure the safety of the priority areas of the Albertine Rift.
Ó2006 Elsevier Ltd. All rights reserved.
1. Introduction
The Albertine Rift is the most species rich region for verte-
brates on the African continent (Brooks et al., 2001; Plumptre
et al., 2003). This part of Africa contains the ‘Mountains of the
Moon’ or Rwenzori Massif that includes Africa’s third highest
peak, the Virunga Volcanoes made famous by its mountain
gorillas, active volcanoes in the Virunga National Park, and
0006-3207/$ - see front matter Ó2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.biocon.2006.08.021
*Corresponding author.
E-mail address: aplumptre@wcs.org (A.J. Plumptre).
BIOLOGICAL CONSERVATION 134 (2007) 178–194
available at www.sciencedirect.com
journal homepage: www.elsevier.com/locate/biocon

Lake Tanganyika – Africa’s deepest lake. The Albertine Rift
encompasses much of the western Rift valley down to south-
ern Tanzania and northern Zambia. We define the region as
extending from 30 km north of Lake Albert to the southern
tip of Lake Tanganyika, including the valley, flanks of the
escarpment and associated protected areas, and the range
of species endemic to it (Plumptre et al., 2003). Various other
publications also recognize the ‘Albertine Rift’ (Poulsen, 1997;
Prigogine, 1985): it is an Endemic Bird Area according to Bird-
Life International (Stattersfield et al., 1998), WWF have de-
fined it as a ‘Global-200’ priority ecoregion (Olson and
Dinerstein, 1998; Burgess et al., 2004), and Conservation Inter-
national (CI) has recognized it as part of the Eastern Afromon-
tane Hotspot in their second global analysis (Brooks et al.,
2004). There is much overlap in the definition of the Albertine
Rift between these publications, but also some differences in
geographical coverage as each analysis has used different cri-
teria to delineate the region. WWF’s Ecoregion focuses pri-
marily on the montane forests and separates moorland
from these forests, while BirdLife International’s Endemic
bird area is broader and includes the moorlands and highland
swamps. CI’s Eastern Afromontane Hotspot is broader still
and includes the definition of the Albertine Rift used here
(see below).
The Albertine Rift is not only important for its biodiversity
but also for its ecological processes and ecosystem services.
The savanna parks contained some of the highest biomasses
of large mammals recorded on earth in the 1960s (Cornet d’Elz-
ius, 1996). War and poaching have led to major decreases in the
numbers of large mammals in these parks but most of the spe-
cies are still present and could recover to former levels with
good protection (Plumptre et al., 2007). The impacts of the
browsing and grazing of the elephants, hippopotamuses, buf-
falos, and antelope species had a major influence on the vege-
tation of the parks (Delvingt, 1978) and as a result of the decline
in numbers of these species it is thought the parks are changing
(Eltringham, 1999). The volcanoes in the Virunga National Park
are active and influence the ecology of a large portion of this
park and its surroundings. The fisheries in some of the lakes
are the most productive on the continent and provide a liveli-
hood for many people (Beadle, 1974; Snoeks, 2000) and the riv-
ers and streams flowing from the forests on the mountains
provide clean water. In Rwanda, for example, it is estimated
that more than 70% of people obtain water that comes from
their national parks (Weber, 1989). The spectacular land forma-
tions and rich biodiversity of the Albertine Rift mean that it has
great potential for tourism. Civil wars and international con-
flict over the past 30 years have hampered tourism develop-
ment but when peace comes to the region there is enormous
potential to develop world class tourism.
This paper summarises the existing literature on biodiver-
sity surveys within sites in the Albertine Rift. It is the result of
a collaborative effort between many different NGOs, pro-
tected area authorities and museums that have information
about this region. Since 2001 the protected area authorities
and their NGO partners have been developing a strategic
framework for conservation in the Albertine Rift. At this time
a core planning group was established, which developed the
strategic framework for conservation in the Albertine Rift,
that includes the Albertine Rift Conservation Society, Dian
Fossey Gorilla Fund International, Institute for Tropical Forest
Conservation, International Gorilla Conservation Programme,
Makerere University Institute of Environment and Natural Re-
sources, Wildlife Conservation Society (WCS) and WWF. The
compilation of the biodiversity information was one aspect
of this planning process and was led by WCS. This paper ad-
vances this collaborative effort by using the collated data to
prioritise sites in the Albertine Rift for conservation according
to their numbers of endemic and globally threatened species.
1.1. The Albertine Rift region
As part of this planning process a series of meetings were
held between February 2001 and March 2003, when the frame-
work was finally developed. During this process it was agreed
that the definition of the Albertine Rift should be as inclusive
as possible for the moment so that over time it could be re-
fined (Plumptre et al., 2003). The current adopted definition
therefore includes all the natural habitats within 100 km east
of the border of Democratic Republic of Congo (DRC) and fol-
lows the 900 m contour line in eastern DRC, including the pro-
tected areas in northern Zambia (Fig. 1). The 900 m contour
was selected because there are museum collections at the
Royal Museum for Central Africa in Tervuren, Belgium, of Alb-
ertine Rift endemic bird species that were found as low as this
altitude.
The total area encompassed is around 313,000 km
2
(Plumptre et al., 2003). The habitats range from the glaciers
and rock at the top of the Rwenzori mountains (5100 m),
down through alpine moorland (3400–4500 m), Giant Senecio
Fig. 1 – Map of the extent of the Albertine Rift as considered
here. Forested protected areas (or surveyed areas) are in
dark grey, savanna/miombo woodland protected areas are
in lighter grey.
BIOLOGICAL CONSERVATION 134 (2007) 178–194 179

and Lobelia vegetation (3100–3600 m), giant heather (3000–
3500 m), raised bogs (3000–4000 m), bamboo forest (2500–
3000 m), montane forest (1500–2500 m), to lowland forest
(600–1500 m), savanna woodland (600–2500 m) and savanna
grassland (600–2500 m). Papyrus and Carex wetlands, together
with lakes and streams, have their own unique habitat types
varying from the rocky and sandy edges to the benthic and
bathypelagic zones in the depths of the lakes. Several very
specialised habitats also occur as a result of the volcanic
activity in the Virunga National Park, including lava flows
and their associated colonising vegetation, hot springs and
species adapted to carbon monoxide and methane.
2. Methods
2.1. Biodiversity of the Albertine Rift
Initially WCS compiled lists of species from existing literature
and by working with researchers at Makerere University,
Uganda. Much of the literature was in unpublished reports
or ‘grey literature’. Data on mammals, birds, reptiles, amphib-
ians, butterflies and plants were compiled, and information
on some museum specimens collated with the help of mu-
seum experts (Royal Museum for Central Africa, Tervuren,
Belgium; Missouri Botanical Gardens, USA; Field Museum,
Chicago, USA, and Royal Botanic Gardens, Kew, UK) from 38
sites within the Albertine Rift (Fig. 2). We also reviewed the lit-
erature for fish species totals as it was recognized that the
lakes in the Albertine Rift region are also important for con-
servation. We attempted to compile information from as
many areas of the Albertine Rift as possible. Data on collec-
tions of amphibians and reptiles from the Royal Museum
for Central Africa allowed some analysis to be made of spe-
cies outside protected areas. Data from plant family descrip-
tions and herbaria were also used to generate a preliminary
list of endemic plant species, many of which are not found
in the sites selected here. Not surprisingly, however, most of
the sites selected were protected areas as that is where most
surveys have been made in the Albertine Rift. However, some
unprotected areas also had survey information, including the
Itombwe Massif, Marungu Massif, high altitude areas west of
Lake Edward and Mt Kabobo as well as areas around Mahale
Mountains National Park. While not covering thewhole Alber-
tine Rift, these sites do include the larger protected areas and
the largest areas of unprotected habitat.
The main sources used for each taxon were as follows:
2.1.1. Mammals
The main source for Uganda was the checklist of mammals of
the national parks (Wilson, 1995) and the small mammal sur-
veys undertaken by the Uganda Forest Department (Howard
and Davenport, 1996). Additional data for large mammals
came from Kingdon (1971–1983) and for small mammals from
Kerbis Peterhans et al. (1996), Kerbis Peterhans and Austin
(1996), Kerbis Peterhans (1997) and van der Straeten and Ker-
bis Peterhans (1999). Dowsett (1990) produced a list of mam-
mals for the Nyungwe Forest Reserve in Rwanda and this
was combined with unpublished sightings from the Wildlife
Conservation Society project in this forest and from Hutterer
et al. (1987). A list for the Virunga Volcanoes was produced
using de Witte (1938), Wilson (1995) and Hutterer et al.
(1987). For DRC, de Witte (1938) reported on extensive surveys
by Belgian scientists in the Virunga National Park. Muhlen-
berg et al. (undated) provided a list for Kahuzi Biega National
Park and Omari et al. (1999) provided a list for Itombwe Mas-
sif. For Burundi, a list was obtained for Kibira National Park
(Field Museum, Chicago (Kerbis Peterhans, in litt., FMNH data-
base, Chicago), Peace Corps and INECN (undated). For Tanza-
nia, lists were obtained for Gombe from the Gombe National
Park website, for Mahale from Anonymous (1985) and for Mbizi
from D. Moyer and W. Stanley’s surveys. For Zambia, lists for
Sumbu and Mweru-Wantipa National Parks were generated
from the distribution records given by Ansell (1978).
2.1.2. Birds
The main source for Uganda was the checklist of birds of the
national parks (Wilson, 1995) and the biological surveys
undertaken by the Uganda Forest Department (Howard and
Davenport, 1996 – which also compiled data from many pre-
viously published sources). Additional data came from the
Enhancement of Research Capacity (ENRECA) project man-
aged by Makerere University Institute of Environment and
Natural Resources, surveys undertaken by WCS, and records
compiled by Malcolm Wilson. Additional published records
that were incorporated after expert peer review include Evans
and Balmford (1992), Gnoske and Marks (1997), Kalina and
Butynski (1996), Friedmann and Williams (1970), Dehn and
Christiansen (2001), Stubblefield (1993), and Allan (1994). For
Rwanda, Kunkel and Kunkel (1969), Dowsett et al. (undated),
Dowsett (1990) and Plumptre et al. (2002) produced lists of
birds for the Nyungwe Forest Reserve. These data were com-
bined with unpublished sightings from the WCS project in
this forest. A list for the Virunga Volcanoes was produced
using Schouteden (1938), Wilson (1982), and Wilson (1995).
Schouteden (1938) reported on extensive surveys by Belgian
scientists in the Virunga National Park in DRC and Verheyen
(1947) added species to the northern part of the park. M. Lan-
guy provided additional observations. Muhlenberg et al. (un-
dated) and Wilson and Catsis (1990) provided a preliminary
list for Kahuzi Biega National Park and this was augmented
by M. Herremans using the database of bird specimens at
the Royal Museum for Central Africa in Tervuren. Schouteden
(1949) surveyed Katanga district from which a list for Mar-
ungu Massif was derived, Prigogine (1960) provided a list for
Mt Kabobo, Prigogine (1971–1984), Wilson and Catsis (1990)
and Omari et al. (1999) provided lists for Itombwe Massif
and Prigogine also provided lists for Idjwi Island (Prigogine,
1967) and for the area west of Lake Edward (Prigogine, 1953).
Many new records in eastern DRC have been published by De-
mey et al. (2000). Tom Butynski contributed records for Mt
Tshiaberimu. Schouteden (1966) published a list of birds of
Burundi with locations where they had been sighted – this
publication was used to compile a list for Ruzizi National
Park. Gaugris et al. (1981) added species to this list and INECN
produced a list for Kibira National Park in the mid 1980 s (IN-
ECN, undated). van de Weghe and Loiselle (1987) also pro-
duced a list for Bururi forest reserve. Neil Baker provided
lists for Gombe Stream and Mahale Mountains National Parks
180 BIOLOGICAL CONSERVATION 134 (2007) 178–194

from the Tanzania Bird Atlas database. These were used to
correct and add to lists compiled by Stanford and Msuya
(1995), Ulfstrand and Lamprey (1960), and Moreau (1943). D.
Moyer provided lists for Mbizi forest.
2.1.3. Reptiles
Pitman (1938, 1974) was used as a starting point for reptiles in
Uganda. Drewes and Vindum (1998) provided a species list for
Bwindi Impenetrable National Park and Vonesh (1998) put to-
gether a list for Kibale National Park. Spawls et al. (2002) iden-
tify localities for species for East Africa using general maps
and these were used to assign species to a site if this was
either mentioned in the text or if the map distribution was
unequivocal. M. Behangana, from recent surveys, provided
several records. Hinkel and Fischer (1988) was used to develop
a list of species for Virunga volcanoes and Nyungwe forest in
Rwanda. This was augmented by de Witte (1941) for the Viru-
nga volcanoes. Dowsett (1990) also provided a list for
Fig. 2 – The northern (a), central (b) and southern (c) portions of the Albertine Rift showing the locations of the various
protected areas (NP = national park; FR = forest reserve; WR = wildlife reserve) or ungazetted areas with species data (no
suffix). Darker shaded areas are forested and lighter areas are savanna grassland or woodland.
BIOLOGICAL CONSERVATION 134 (2007) 178–194 181

Nyungwe Forest. D. Meirte extracted lists of specimen loca-
tions from the database at the Royal Museum for Central Afri-
ca in Tervuren. For DRC, de Witte (1941) produced a list of
reptiles for Virunga National Park and mapped the distribu-
tions of chameleons in central Africa (de Witte, 1965). D. Mei-
rte extracted lists of specimen locations from the database at
the Royal Museum of Central Africa in Tervuren. Only limited
data were obtained for Burundi. Spawls et al. (2002) was used
to compile a list of reptiles for Gombe and Mahale Mountains
Parks in Tanzania.
2.1.4. Amphibians
Drewes and Vindum (1994, 1998) and Drewes et al. (1992) pro-
vided a species list for Bwindi Impenetrable National Park
and Vonesh (1998) put together a list for Kibale National Park
in Uganda. Many recordswere provided by M. Behangana from
recent surveys. In Rwanda, Hinkel and Fischer (1988) was used
to develop a list of species for Virungavolcanoes and Nyungwe
forest. This was augmented by de Witte (1941) for the Virunga
volcanoes. Dowsett (1990) also provided a list for Nyungwe For-
est. For DRC, de Witte (1941) produced a list of amphibians for
Virunga National Park but the identifications needed updating.
The database at the Africa Museum in Tervuren is in the pro-
cess of being updated for amphibians to reflect current taxon-
omy and we used the corrected data in this database to create
lists of species for DRC and to add records to sites. Laurent
(1972) increased the amphibian list of de Witte for Virunga Na-
tional Park and corrected some mis-identifications. Laurent
(1964) was used to compile a list of amphibians for the Itombwe
Massif. D. Meirte provided corrections to the taxonomy of the
older literature and also added many species from the database
at the Royal Museum for Central Africa at Tervuren. Some data
on distribution and systematics of tree frogs were taken from
Schiøtz (1999). Only limited data were obtained for Burundi
and none for Tanzania.
2.1.5. Butterflies
Information was drawn from a variety of sources including Car-
casson (1961, 1975), d’Abrera (1980, 1997), Henning (1988), Kiel-
land (1990), Larsen (1991), Ackery et al. (1995), Davenport (1996),
Howard and Davenport (1996), Congdon and Collins (1999),
Congdon et al. (2001), as well as numerousworkers from earlier
parts of the last century referenced in these publications. Addi-
tional information came from collections held at Makerere
University Zoology Museum, Kampala, and the National Muse-
ums of Kenya, Nairobi. Steve Collins (ABRI, Nairobi) provided
considerable and invaluable information, and we are very
grateful to C. Congdon (Tanzania) and A. Gardiner (Zambia)
for very useful comments on an earlier draft.
2.1.6. Plants
The main starting point for Uganda was the tree surveys
undertaken by the Uganda Forest Department (Howard and
Davenport, 1996). Additional data was added by G. Eilu (climb-
ers) Poulsen (1997, terrestrial herbs) and D. Hafashimana (epi-
phytes) for several forests. WCS has also been surveying
many of the Rift forests over the past year and the species
identified were incorporated in the database by D. Nkuutu.
Lock (1977) provided a list of species for Queen Elizabeth Na-
tional Park. Nabanyumya (1991) listed trees for Kalinzu and
Maramagambo forests. Synnott (1985) provided a checklist
of plants for Budongo Forest Reserve. For Rwanda, plant spe-
cies lists for the Virunga Volcanoes were obtained from Burtt
(1934), Robyns (1948–1955), Troupin (1978–1988), and the her-
barium at the Karisoke Research Station. Plant species for
Nyungwe were compiled from Robyns (1948–1955), Troupin
(1992), Plumptre et al. (2002) and the herbarium at the Project
Conservation de la Foreˆt de Nyungwe. Robyns (1948–1955)
provided a relatively complete list for Virunga National Park
in DRC and Fischer (1996) provided a list for Kahuzi Biega Na-
tional Park. No plant lists were obtained for sites in Burundi.
For Tanzania, the list of plants of Gombe was provided by Roy
Gereau from his surveys there. Toshisada Nishida kindly pro-
vided a list of plants eaten by chimpanzees for Mahale Moun-
tains National Park. Additional species for Mahale were
obtained from Vollesen and Bidgood (1996, 1999). A species
list for Mbizi forest was compiled from Mwasumbi (2000).Leb-
run and Stork (1991–1997) was used to correct synonyms to a
standardised list of names.
2.1.7. Data cleaning and identification of species of
conservation concern
After six months of developing draft lists a meeting was con-
vened by the core planning group to bring together experts
from the region, particularly from museums and other insti-
tutions to review and augment the draft species lists. This
meeting also addressed threats to these sites and refined
the lists of endemic species for the Albertine Rift. Following
this meeting there was another period of six months adding
to the data and in particular checking species names and syn-
onyms because taxonomy of many species has changed over
the 70 years of publications that were used. WCS and Make-
rere University were involved in this data cleaning.
Numbers of endemic and globally threatened species were
calculated for each site and used to prioritise sites in the Alb-
ertine Rift for conservation as follows. Endemic species lists
were compiled with the help of experts in each taxonomic
field. The mammal list came from Conservation International
(J. Pilgrim) with additions by J. Kerbis Peterhans. The list com-
piled by Stattersfield et al. (1998) was used for birds. This pub-
lication recognized two areas of endemism in this region; the
Albertine Rift and the Eastern Zairean Region of endemism.
However, some species endemic to the Albertine Rift overlap
substantially with those from the Eastern Zairean region (M.
Herremans – data from Africa Museum at Tervuren) and the
planning group made the decision that it was better to com-
bine these two areas in this analysis. Lists of endemic reptile
and amphibian species were compiled by D. Meirte. T. Daven-
port compiled a list of endemic butterfly species in collabora-
tion with S. Collins, C. Congdon and A. Gardiner. E. Ndomba, P.
Ssegawa, G. Eilu and A. Plumptre worked with botanists at the
Royal Botanic Gardens, Kew, and Missouri Botanical Gardens
to develop a preliminary list of endemic plant species based
upon published family descriptions from most existing Afri-
can floras (East Africa, Congo, Cameroon, Gabon, and Zambe-
siaca). This list is likely to miss many species because many
families have not been described in DRC or East Africa. Ende-
mic species lists are given in Plumptre et al. (2003) and are
also at www.albertinerift.org. Globally threatened species
were derived from the 2002 IUCN Red List (Hilton-Taylor,
182 BIOLOGICAL CONSERVATION 134 (2007) 178–194

2000 and updated lists on the associated website www.iucn-
redlist.org) and BirdLife International (2000).
2.2. Ranking sites for conservation
While there is a danger that ranking sites in terms of their
conservation value can lead to the lower-ranked sites being
sidelined, it is still important that priorities are identified
given the limited human and financial resources available
for conservation. However, even the poorest sites for which
we have compiled data are rich, in a global context, have
many restricted-range species, and therefore deserve conser-
vation attention.
Ranking using species data requires subjective decisions,
particularly when data are incomplete. Only five taxa (mam-
mals, birds, reptiles, amphibians and plants) can be used in this
study of the Rift because species data are not collated or avail-
able for other taxa. In the absence of more extensive data, we
can only hope that these taxa act as good surrogates for others.
Several aspects of surrogacy were tested between these taxa.
This indicated that, in many cases, one taxon could act as a rea-
sonable surrogate for the other four across the sites surveyed
(Plumptre et al., 2003). Numbers of endemic species for one tax-
on correlated well (Pearson Correlations) with numbers of en-
demic species in another taxon, for all possible combinations
(P< 0.001 for all tests). Surrogacy for threatened species was
not so good. Numbers of threatened mammals, amphibians
and birds correlated well between sites but threatened plants
and reptiles did not show any significant correlation with other
taxa. However, plants and reptiles have been less completely
assessed for threatened species than the other three taxa. Only
two reptiles were classified as threatened in this region in the
2002 Red List. If total numbers of species are used to rank sites
then plants will dominate the rankings because of the larger
number of species. The general public may wish to rank sites
on mammal and bird fauna because these are popular. Alterna-
tively, it might be better to prioritise sites on species that have
economic value and can attract tourists or provide livelihood to
local communities, or by their ecological role in the ecosystem.
Weightings could be made for certain species or certain taxa to
incorporate these ideas. Here we have decided to weight taxa
equally so that a high number of mammal species at a site re-
ceives the same rank as a high number of plants irrespective
of the actual number of species. This effectively gives equal
weight to any taxon and ranks biodiversity per se rather than so-
cial, economic or ecological functions of biodiversity.
Sites were ranked in terms of the number of endemic and
number of globally threatened species for each taxon. These
ranking scores were standardized for each site by dividing by
the maximum rank score to account for the fact that some sites
do not have data for all taxa and to account for the varying
number of endemic or threatened species between taxa. For
example, only two reptile speciesare currently listed as threa-
tened of the 175 species found in the Albertine Rift sites (pri-
marily because reptiles have not been well assessed). Three
sites have threatened reptiles, which means that all sites with
no threatened reptiles scored a value of ‘4’ in the rankings.
However a rank value of ‘4’ for birds is a high value with many
threatened species and therefore ranks cannot be summed
across taxa without standardizing them by dividing by the
maximum rank (4 in the case of reptiles).A mean rank was then
calculated for both endemic and globally threatened species
for each site across the five taxa by summing ranks from each
taxon and dividing this by the number of taxa for which there
were survey data (i.e., for which an initial rank was even possi-
ble). These final standardised scores were then ranked (1–38) to
rank all the available sites for which we compiled data. Thus,
the final rankings allow better comparison of siteswith varying
levels of knowledge. In presenting the results we grouped sites
into categories of high, medium and low rank rather than using
actual ranking values for each site. This avoids false precision
because the data for many sites are incomplete, some mean
ranking scores are based upon just one taxon, and the effort
made in sampling sites varies widely. Comparing the rankings
for endemic and globally threatened species allows an analysis
of the relative importance of these two criteria.
The ranking process described above did not attempt to
correct for the area of the sites. We were more interested in
which are the priority sites for species conservation in the
Albertine Rift rather than which are the richer sites relatively
when area is standardised. From a conservation perspective
the larger a site is, the easier it is to manage compared to
many smaller sites of the same total area; conserving a few
large sites will in general be the cheaper option. However,
for comparison with other studies we did make this correc-
tion. We corrected for area by calculating
scored endemic or threatened species
¼lnðnumber of species þ1Þ=lnðareaÞ:
3. Results
The results show that this region contains more than half of
continental Africa’s bird species and nearly 40% of its mammal
species. Reptile and amphibian species do not appear as abun-
dant but this may be a function of the effort that has been
made in collecting, identifying and cataloguing them in this re-
gion. While invertebrate taxa have been poorly surveyed, this
region is known to have a large number of endemic butterflies
(probably the best surveyed invertebrate taxon). The results
presented here are the current state of knowledge, which will
be improved on in due course as more surveys are conducted.
3.1. Biodiversity values
3.1.1. Mammals
No endemic families occur in the Albertine Rift but two ende-
mic genera occur, Rwenzorisorex, and Delanymys. A total of 402
mammal species (158 genera and 46 families) have been re-
corded in the Albertine Rift, of which 35 are endemic (Tabl e 1 ).
Most of the endemic mammals are shrews and rodents. Of
the larger endemic mammals, the eastern gorilla is the best
known and has two subspecies (mountain, Gorilla beringei berin-
gei, and Grauer’s, Gorilla beringei graueri, gorillas). The Rwenzori
duiker, Cephalophus rubidus, and the golden monkey, Cercopithe-
cus kandti, are two other medium sized endemic mammals.
Small mammals have been poorly surveyed throughout much
of the Rift, particularly towards the southernend, and it is very
likely more species would be added with further effort.
BIOLOGICAL CONSERVATION 134 (2007) 178–194 183

Virunga National Park in eastern DRC has the highest
number of endemic mammal species (21). Bwindi Impenetra-
ble National Park (20) and Rwenzori Mountains (18) rank next
highest.
Thirty-four mammal species are globally threatened (Crit-
ically Endangered, Endangered or Vulnerable) according to
the 2002 IUCN Red List (Hilton-Taylor, 2000;www.iucnred-
list.org), of which 12 are Albertine Rift endemics. These in-
clude eastern gorilla (Gorilla beringei), golden monkey and
Rwenzori otter shrew Micropotamogale ruwenzorii. Kahuzi
Biega National Park had the highest number of globally
threatened mammals (14) followed by Virunga National Park
(13).
3.1.2. Birds
Three genera are endemic to this region, Pseudocalyptomena,
Graueria, and Hemitesia but no families are endemic. At least
1061 bird species (in 368 genera and 80 families) occur in
the Albertine Rift (Table 2) of which 4.5% are migratory spe-
cies that overwinter in the region but do not breed, or which
pass through on migrations within the African continent.
Although this is the most thoroughly surveyed group of ani-
mals, new species for the Rift continue to be added as migrant
species and new range extensions are recorded. Within our
definition of the Albertine Rift there are two contiguous Ende-
mic Bird Areas (EBAs) defined by BirdLife International: Alber-
tine Rift and Eastern Zairean Lowlands (Tab l e 2)(Stattersfield
et al., 1998). We have combined these areas because old mu-
seum collections of Albertine Rift endemic species show they
occur at lower altitudes in eastern DRC and that they overlap
in altitudinal range with the Eastern Zairean lowland species
(Bober et al., 2001; Herremans et al., 2002). The total number
of endemic birds in the Albertine Rift as defined here is 41
species, which includes the endemics of these two EBAs.
The Itombwe Massif, an unprotected area west of the
northern end of Lake Tanganyika, contains more endemic
species than any other site in the Albertine Rift (34). It is clo-
sely followed by Kahuzi Biega National Park (30) and Virunga
National Park (27).
Twenty-five Albertine Rift species are globally threatened,
13 of which are endemics. These include congo bay owl
(Phodilus prigoginei), itombwe nightjar (Caprimulgus prigoginei),
Table 1 – Richness of mammals across sites of the Albertine Rift Mountains, including number of species,
number of Albertine Rift (AR) endemic species, and number of globally threatened species (CR = Critically Endangered,
EN = Endangered, VU = Vulnerable)
Site Species no. AR endemic species Threatened CR, EN, VU
Budongo FR
a
95 0 5
Bugoma FR 38 0 4
Bugungu WR 9 0 1
Bururi FR 9 1 1
Bwindi Impenetrable NP
a
135 20 7
Echuya FR
a
24 7 1
Forest West of Lake Edward 8 0 1
Gombe NP 19 1 4
Ibambaro FR 2 0 0
Itombwe Massif 72 4 10
Itwara FR 18 0 0
Kagombe FR 14 0 3
Kahuzi Biega NP
a
136 15 14
Kalinzu–Maramagambo FR
a
58 1 3
Karuma WR 57 0 4
Kasyoha–Kitomi FR
a
47 2 3
Kibale NP
a
115 5 7
Kibira NP
a
71 8 7
Kitechura FR 17 0 1
Kyambura WR 37 0 3
Mafuga FR 20 3 1
Mahale Mountains NP
a
52 1 6
Matiri FR 12 1 0
Mbizi FR 23 1 2
Murchison Falls NP
a
109 0 5
Mweru-Wantipa NP
a
50 0 7
Nyungwe NP
a
86 14 3
Queen Elizabeth NP
a
97 0 6
Rwenzori Mountains NP
a
102 18 10
Semliki NP
a
86 1 5
Semliki WR 69 0 4
Sumbu NP
a
61 0 6
Virunga NP
a
196 21 13
Total 402 35 36
a Reasonably surveyed for all mammals.
184 BIOLOGICAL CONSERVATION 134 (2007) 178–194

kungwe apalis (Apalis argentea), grauer’s rush warbler (Brad-
ypterus graueri) and golden-naped weaver (Ploceus aureonucha).
Itombwe had the highest number of globally threatened spe-
cies (15) followed by Virunga National Park (11), Kahuzi Biega
National Park (11) and the mountains west of Lake Edward
(11).
3.1.3. Reptiles
No endemic genera or families of reptiles are known from the
Albertine Rift. A total of 175 reptiles, from 69 genera and 20
families (about 14% of Africa’s reptiles) have been recorded
for the Albertine Rift (Tabl e 3). Far fewer sites have been sur-
veyed for reptiles to the same extent as for mammals and
birds, although some records existed for at least 33 sites.
Itombwe Massif and Kahuzi Biega National Park in eastern
DRC may contain many species but to date have been poorly
surveyed. Other areas that need work include the Marungu
Massif in eastern DRC at the southern end of Lake Tanganyika
and the Mahale Mountains National Park and its surrounding
natural vegetation.
Sixteen endemic reptile species occur in the Rift, of which
Virunga National Park contains the highest number (11), fol-
lowed by Rwenzori Mountains National Park (9) and Nyungwe
National Park (8). Endemic species include five chameleons
such as the strange-horned chameleon (Bradypodion xenorhi-
num), and Johnston’s chameleon (Chamaeleo johnstoni), two
colubrid snakes, one viper, six skinks, one worm snake (Lepto-
typhlops) and one lacertid lizard.
Only two globally threatened reptiles are currently listed
for the Albertine Rift (Trionyx triunguis and Osteolaemus tetra-
spis). However, this is because the region has few data and
reptiles as a whole have not been assessed completely for
their threatened status. The IUCN Global Reptile Assessment
is underway and it is likely many more species will be added
to the Red List. As a result no site has more than one globally
threatened reptile (Table 3).
3.1.4. Amphibians
Three amphibian genera are endemic to the Albertine Rift;
Laurentophryne,Chrysobatrachus and Callixalus. There are 119
Table 2 – Richness of birds across sites of the Albertine Rift Mountains, including number of species, number of Albertine
Rift (AR) endemic species as defined by BirdLife International (with number of eastern Zairean lowland endemic species in
parentheses that were included in the definition of Albertine Rift Endemic species here – see text) and number of globally
threatened species (CR = Critically Endangered, EN = Endangered, VU = Vulnerable)
Site Species no. AR endemic species Threatened CR, EN, VU
Budongo FR 362 0 1
Bugoma FR 221 0 1
Bururi FR 155 13 3
Bwindi Impenetrable NP 381 24(1) 6
Echuya FR 136 14 2
Forests West of Lake Edward 420 25 11
Gombe NP 267 0 2
Idjwi 150 2 1
Itombwe Massif 583 34(4) 15
Itwara FR 183 0 0
Kagombe FR 121 0 0
Kahuzi Biega NP 335 32(3) 11
Kalinzu–Maramagambo FR 393 4 1
Kasyoha–Kitomi FR 308 2 1
Kibale NP 327 3 3
Kibira NP 211 21 7
Kitechura FR 90 0 0
Kyambura WR 450 0 6
Lendu Plateau 317 6 4
Mafuga FR 130 10 0
Mahale Mountains NP 250 2 1
Marungu 282 1 0
Matiri FR 119 0 0
Mbizi FR 116 0 0
Mt Kabobo 231 18 3
Murchison Falls NP 476 0 7
Nyungwe NP 280 26 7
Queen Elizabeth 594 0 7
Rusizi NR 182 1 3
Rwenzori Mountains NP 241 21 4
Semliki NP 441 7(5) 9
Semliki WR 435 0 3
Virunga NP 706 27(2) 11
Total 1061 41(6) 25
In the analyses presented in this paper Albertine Rift and Eastern Zairean Lowland endemic species were combined because of the extensive
overlap in distributions of the two groups as can be seen here.
BIOLOGICAL CONSERVATION 134 (2007) 178–194 185

species of amphibians in the Albertine Rift, including 29 gen-
era and 11 families, (about 19% of Africa’s amphibians).
Thirty-six endemic species have been identified. It is likely
that more survey effort would uncover further endemic spe-
cies. Virunga National Park had the highest number of ende-
mic species (16) with Itombwe Massif (16) and Nyungwe Park
(14). Four endemic species (Hyperolius pustulifer,Schoutedenella
loveridge,Schoutedenella mossoensis,Schoutedenella vercammeni),
have only been recorded outside the 40 sites described here
and are thus not found in any of the protected areas in the
Albertine Rift.
Sixteen Albertine Rift amphibians are globally threatened,
of which 14 are endemic. Itombwe Massif has more threa-
tened species (CR, EN or VU) than other sites (11) followed
by Virunga National Park with 10 and Bwindi Impenetrable
National Park with six (Tab le 4).
3.1.5. Fish
While no attempt was made to put together species lists of
fish for the rivers and lakes in the Albertine Rift, the literature
was searched for data on the major lakes (Albert, George, Ed-
ward, Kivu and Tanganyika). Lake Tanganyika alone has 289
endemic species that make up 89% of fish diversity of the lake
(Snoeks, 2000). Only Lake Malawi has more endemic fish in
Africa. Fifty-six fish species are endemic to lakes George
and Edward, while Kivu and Albert have 15 and six endemic
fish respectively. Only 10% of Lake Tanganyika’s shore has
been explored and a total of over 1200 faunal species (verte-
brates and invertebrates) have been recorded, making it the
second highest recorded diversity for any lake on earth (Patt-
erson and Makin, 1998).
3.1.6. Butterflies
The total number of butterfly species found in the Albertine
Rift is unknown, as this information cannot be compiled un-
til many more areas have been surveyed, particularly in
eastern DRC. In Uganda, inventories of the forests in the
Albertine Rift have shown that at least 581 species of butter-
fly, 16% of the estimated 3630 species in Africa, occur in this
part of the Albertine Rift alone (Howard and Davenport,
1996). It is possible that, given the numbers from Uganda
and Tanzania, up to 1300 butterfly species might occur in
the Rift, about 35% of Africa’s total. There are no known en-
demic families, but the genus Kumothales is restricted to the
Albertine Rift. It is known that 117 endemic species from 49
genera exist in the Albertine Rift (Plumptre et al., 2003). The
Table 3 – Richness of reptiles across sites of the Albertine Rift Mountains, including number of species, number
of Albertine Rift (AR) endemic species, and number of globally threatened species (CR = Critically Endangered,
EN = Endangered, VU = Vulnerable)
Site Species no. AR endemic species Threatened CR, EN, VU
Budongo FR 48 1 0
Bugoma FR 9 0 0
Bugungu WR 9 0 0
Bururi FR 1 1 0
Bwindi Impenetrable NP 34 6 0
Echuya FR 4 0 0
Forests West of Lake Edward 6 3 0
Gombe NP 1 0 0
Itombwe Massif 35 5 0
Itwara FR 10 0 0
Kahuzi Biega NP 69 7 0
Kalinzu–Maramagambo FR 9 0 0
Karuma WR 15 0 0
Kasyoha–Kitomi FR 9 0 1
Kibale NP 56 3 0
Kibira NP 3 2 0
Kyambura WR 12 0 0
L. Rukwa 7 0 0
L. Tanganyika 13 0 0
Lendu Plateau 6 0 0
Mafuga FR 17 2 0
Mahale Mountains NP 4 0 0
Marungu Massif 6 0 0
Mbizi FR 3 0 0
Mt Kabobo 6 2 0
Murchison Falls NP 32 0 1
Nyungwe NP 43 8 0
Queen Elizabeth 34 0 0
Rusizi NR 3 0 0
Rwenzori Mountains NP 34 9 0
Semliki NP 49 0 0
Semliki WR 33 0 1
Virunga NP 109 11 0
Total 175 16 2
186 BIOLOGICAL CONSERVATION 134 (2007) 178–194

total number of endemics (of which 85% are forest depen-
dent) is considerably larger than the 78 endemic species
found in the Eastern Arc Mountains and coastal forests of
Tanzania and Kenya, and could increase with further survey
effort. Whether or not butterflies are indicative of other
invertebrate species is unclear but these numbers do dem-
onstrate unequivocally that this region is not only important
for vertebrate conservation.
3.1.7. Plants
Higher plants have been relatively well surveyed in the forests
of Uganda and Rwanda but elsewhere in the Albertine Rift
surveys have been patchy. Currently 5793 plant species (from
1537 genera and 233 families) have been recorded within the
Rift but this will change as surveys are discovering new spe-
cies regularly even within Uganda (Tabl e 5 ). These data in-
clude ferns and higher taxa but do not include the
bryophytes and lichens, which are very poorly surveyed.
The number of plant species is high compared with many re-
gions of similar size and forms 14% of all mainland Africa’s
estimated plant species. A preliminary estimate of the num-
bers of endemic plant species has been compiled by the
WCS’s Albertine Rift Programme and this now numbers 551
species. These lists are based on published flora descriptions
of plant families, but many families have not been described
for DRC or East Africa. Some input was made by herbaria ex-
perts, but the list is still incomplete. As a result this list
should be considered to be very preliminary and could in-
crease greatly when lower plants and little studied growth
forms such as climbers, epiphytes, lichens and bryophytes
are included. Western Tanzania, especially around Mahale
Mountains National Park, appears to be particularly rich in
plant species and yet has still not been surveyed intensively.
As such it deserves more attention. Virunga National Park
in eastern DRC and Bwindi Impenetrable National Park in
Uganda had the highest numbers of plant species recorded
but both sites have been relatively intensively surveyed. The
Marungu Massif and Itombwe Massif in eastern DRC have
few records but also could be relatively rich and require
surveys.
3.2. Site priority rankings
Rankings were made for numbers of both endemic and glob-
ally threatened species for all taxa (mammals, birds, reptiles
amphibians and plants), for each site as explained in Section
2.2 (Tab le 6). We then grouped the sites into high (rank scores
1–12), medium (rank scores 13–24) and low (rank scores 25–38)
scoring sites for both criteria and plotted the results in a two-
way table for endemic and globally threatened species (Table
7). Given the gaps in the data and differences in sampling ef-
fort between sites we believe that grouping the sites into the
three broad ranking categories provides a more conservative
approach by alleviating these biases.
Those six sites that scored highest are considered to be the
most important because they rank highly for number of both
Table 4 – Richness of amphibians across sites of the Albertine Rift Mountains, including number of species, number
of Albertine Rift (AR) endemic species, and number of globally threatened species (CR = Critically Endangered,
EN = Endangered, VU = Vulnerable)
Site Species no. AR endemic species Threatened CR,EN, VU
Budongo FR 32 1 1
Bugoma FR 20 1 0
Bururi FR 4 4 1
Bwindi Impenetrable NP 29 6 6
Echuya FR 19 5 1
Forests West of Lake Edward 6 6 3
Itombwe Massif 23 16 11
Itwara FR 19 0 0
Kahuzi Biega NP 25 7 4
Kalinzu–Maramagambo FR 25 2 2
Karuma WR 16 0 0
Kasyoha–Kitomi FR 16 3 2
Kibale NP 33 5 3
Kibira NP 1 0 0
Kitechura FR 15 0 0
Kyambura WR 14 0 0
Mafuga FR 1 1 0
Marungu 19 1 0
Matiri FR 15 0 0
Mt Kabobo 8 7 5
Murchison Falls NP 14 0 0
Nyungwe NP 33 14 5
Queen Elizabeth NP 10 1 1
Rwenzori Mountains NP 25 7 1
Semliki NP 24 1 0
Semliki WR 13 0 0
Virunga NP 65 16 10
Total 119 36 16
BIOLOGICAL CONSERVATION 134 (2007) 178–194 187

endemic and globally threatened species. Virunga National
Park consistently ranks high for all taxa because it contains
a very diverse suite of habitats ranging from glaciers to low-
land forest and savannas. Itombwe Massif is a critical area
for conservation as it is currently unprotected and yet ranks
in the top five for endemic and globally threatened species.
Kahuzi Biega National Park in eastern DRC is just to the north
of Itombwe and ranks highly despite being poorly surveyed.
Two sites in Uganda rank highly: Bwindi Impenetrable Na-
tional Park and Kibale National Park. These two sites include
forest at high and medium altitude respectively. Nyungwe
National Park ranks highly although the contiguous Kibira
National Park does not rank so highly, probably because it
has been less surveyed. The next important sites are those
three that rank highly for globally threatened species and
medium for endemic species richness, as globally threatened
species are in more urgent need of conservation. On the
whole this classification seems to make sense intuitively
and from what is known about these sites.
These rankings are affected by area. Those sites that are
large tend to have more endemic species and a higher species
richness. In some studies a correction is made for the area of
the site to calculate those sites that have the highest numbers
of species per unit area. We do not believe this is as useful for
conservation purposes because we are ideally trying to con-
serve the largest sites with most endemic and globally threa-
tened species. However, it is useful for comparisons with
other studies to calculate those sites that have high numbers
of endemic and globally threatened species per unit area. The
results show that many of the key sites previously identified
still rank highly (Tabl e 8 ). Large sites such as Virunga and Ka-
huzi Biega National Parks and the Itombwe Massif contain
large numbers of endemic and globally threatened species
per unit area as well as in total.
4. Discussion
The Albertine Rift contains many high global conservation
priority sites. This region contains more vertebrate and more
endemic vertebrate species than anywhere else on the Afri-
can continent (Burgess et al., 2004). Although for many taxa
and sites, species lists are still incomplete and will increase
as more research is undertaken, the data presented here do
show the large number of species known from this region.
The data are used to prioritise sites for conservation but we
also caution how this ranking is interpreted and used. Most
surveys have focused on the protected areas listed here and
yet very little is known about the surrounding landscapes in
which these sites sit. Given this low level of knowledge out-
side protected areas it makes sense to also conserve at a lar-
ger landscape scale until we have a better knowledge of what
occurs elsewhere. Managing at the landscape scale in the Rift
is a necessary long-term conservation strategy, even though it
will require more resources than focusing on single protected
areas. Management at a landscape scale will also ensure that
certain species, landscape species (Sanderson et al., 2002),
may stand a better chance of survival over the longer term
(Plumptre et al., 2007). For example, large predators such as
leopards Panthera pardus, lions Panthera leo, some of the
larger primates (chimpanzees Pan troglodytes, gorillas Gorilla
Table 5 – Richness of plants across sites of the Albertine Rift Mountains, including number of species, number of tree
species only, number of Albertine Rift (AR) endemic species, and number of globally threatened species (CR = Critically
Endangered, EN = Endangered, VU = Vulnerable)
Site Species no. No. tree species AR endemic species Threatened CR, EN, VU
Budongo FR
a
1064 449 29 18
Bugoma FR 256 245 7 12
Bwindi Impenetrable NP
a
1405 393 74 18
Echuya FR
a
423 131 32 1
Gombe NP
a
510 112 12 0
Itwara FR 258 248 7 10
Kagombe FR 211 201 3 5
Kahuzi Biega NP
a
1171 218 145 9
Kalinzu–Maramagambo FR
a
787 442 34 12
Kasyoha–Kitomi FR
a
901 419 41 17
Kibale NP
a
532 330 16 12
Kitechura FR 113 108 2 0
Mafuga FR 115 100 7 2
Mahale Mountains NP
a
1174 220 39 9
Matiri FR 113 105 2 2
Mbizi FR
a
385 94 18 8
Murchison Falls NP 149 145 1 5
Nyungwe Forest
a
1105 230 137 7
Queen Elizabeth
a
950 288 22 5
Rwenzori Mountains NP
a
696 199 55 5
Semliki NP 333 318 7 14
Virunga NP
a
2077 264 230 10
Total 5793 821 551 40
a Reasonably surveyed for all plant groups (ferns, herbs, climbers and shrubs).
188 BIOLOGICAL CONSERVATION 134 (2007) 178–194

beringei), and large ungulates (elephants Loxodonta africana,
hippopotamuses Hippopotamus amphibious) are species that
need large areas to maintain viable populations. As such
some of the ‘less rich’ sites and as yet unsurveyed ones
may have important connectivity roles and should not be
ignored.
4.1. Contiguous sites and conservation
Many of the protected areas or conservation sites in the Alb-
ertine Rift are contiguous with other protected areas/sites or
are still connected by relatively natural habitat. Where these
connections are truncated existing biota can be seriously
jeopardised (e.g., Kibale-QENP Corridor, Kahuzi Biega low-
land–highland sectors, Kibira (Teza sector in the south) and
its connection to Nyungwe National Park). These natural hab-
itats serve as corridors for wildlife, and their conservation
status is in most cases unclear because they have been little
surveyed. As such they form larger ‘landscapes’ whose spe-
cies richness will be larger than for single sites. Larger conser-
vation areas have higher chances of long-term persistence of
their species and habitats (Groves, 2003). Many of these land-
scapes cross international boundaries or connect sites that
are managed by different institutions, such as the forest re-
serves and national parks in Uganda. If protected areas are
to persist it is important that they are managed as one contig-
uous unit rather than independent sites in order to maintain
connectivity. Transboundary conservation and inter-institu-
tional management of larger landscapes is complex and re-
quires regular coordination if it is to work (see Plumptre
et al., 2007). However for an area such as the Albertine Rift,
with heavy human population pressure, many conservation
sites are becoming islands and maintaining connectivity is a
priority.
Where do these landscapes occur? The largest and most
critical of the landscapes includes the Virunga National Park
in DRC, with the Parc National des Volcans in Rwanda, and
Table 6 – Rankings for each taxon at each site for endemic (End) and globally threatened (Th) species
Site Mamm End Mamm Th Bird
End
Bird Th Rep End Rep Th Amph
End
Amph Th Plant End Plant Th
Budongo FR 19 12 21 20 12 4 14 11 10 1
Bugoma FR 19 15 21 20 14 4 14 16 15 5
Bugungu WR 19 25 14 4
Bururi FR 12 25 11 14 14 4 11 11
Bwindi Impenetrable NP 2 5 6 10 5 4 7 3 4 1
Echuya FR 7 25 10 18 14 4 9 11 9 20
Forest West of Lake Edward 19 25 5 2 7 4 7 7
Gombe NP 12 15 21 18 14 4 14 21
Ibambaro FR 19 31
Idjwi 0 17 20 9 4
Itombwe Massif 9 3 1 1 14 4 1 1
Itwara FR 19 31 21 26 14 4 20 16 15 8
Kagombe FR 19 19 21 26 19 13
Kahuzi Biega NP 4 1 2 2 4 4 4 6 2 10
Kalinzu–Maramagambo FR 12 19 15 20 14 4 13 9 8 5
Karuma WR 19 15 14 4 20 16
Kasyoha–Kitomi FR 11 19 17 20 14 1 12 9 6 3
Kibale NP 8 5 16 14 7 4 9 7 13 5
Kibira NP 6 5 7 6 12 4 20 16
Kitechura FR 19 25 21 26 20 16 20 21
Kyambura WR 19 19 21 10 14 4 20 16
Lendu Plateau 14 12 14 4
Mafuga FR 10 25 12 26 9 4 14 16 15 18
Mahale Mountains NP 12 9 17 20 9 4 7 10
Marungu 20 26 14 4 14 16 13
Matiri FR 12 31 21 26 20 16 20 18
Mbizi FR 12 24 21 26 14 4 12 13
Mt Kabobo 9 14 14 4 4 4
Murchison Falls NP 19 12 21 6 14 1 20 16 22 13
Mweru-Wantipa 19 5
Nyungwe NP 5 19 4 6 3 4 3 4 3 12
Queen Elizabeth NP 19 9 21 6 14 4 14 11 11 13
Rusizi NR 64
Rwenzori Mountains NP 3 3 7 12 2 4 4 11 5 13
Semliki NP 12 12 13 5 14 4 14 16 15 4
Semliki WR 19 15 21 14 14 1 20 16
Sumbu 19 9
Virunga NP 1 2 3 2 1 4 1 2 1 8
The lowest numbers are the sites with most species. These ranks were then standardized by dividing each rank value by the highest number in
each column. Mamm – mammal; Rep – Reptile; Amph – amphibian.
BIOLOGICAL CONSERVATION 134 (2007) 178–194 189

Semliki, Rwenzori, Bwindi Impenetrable, Queen Elizabeth,
and Kibale National Parks and Kasyoha–Kitomi and Kalin-
zu–Maramagambo Forest Reserves and Kigezi and Kyambura
Wildlife Reserves in Uganda. This ‘Greater Virunga landscape’
covers about 13,190 km
2
and includes a wide variety of habi-
tats and altitudes, ranging from 600 to 5100 m above sea level.
It is also incredibly rich in total species as well as endemic
and globally threatened species (Table 9,Plumptre et al.,
2007) and is one of the most biodiverse sites in the world.
There is nowhere else in Africa that can claim vertebrate spe-
cies numbers close to those found here and detailed studies
of sites in the neotropics certainly have fewer numbers than
these (Gentry, 1990). Despite the large size of this landscape,
there are species that still occur at low density so there is a
need to manage it as a whole (Plumptre et al., 2007). However
as all these protected areas are already linked, there is only a
need to generate the political will to manage this landscape
as one protected area across the international boundaries.
The Lendu Plateau (Blue Mtns) to the north may also be an
important addition to this landscape. It is the peripheral for-
ests that may contain biological and genetic outliers due to
their relatively longer term isolation.
Table 8 – Relative rankings for numbers of endemic and globally threatened species when area is standardised
AR endemic species Globally threatened species
High Medium Low
High Bwindi NP
Nyungwe NP
Virunga NP
Itombwe Massif Echuya FR
Kahuzi Biega NP Bururi FR Mafuga FR
Mt Kabobo Rusizi NR
Rwenzori NP
Forest W. of L. Edward
Medium Kasyoha–Kitomi FR Budongo FR
Kibira NP Kalinzu–Maramagambo FR Idjwi
Kibale NP Lendu Plateau Gombe NP
Semuliki NP Queen Elizabeth NP
Mahale NP
Mbizi FR
Low Kagombe FR Marungu Massif
Semliki WR Bugungu WR
Kyambura WR Ibambaro FR
Bugoma FR Karuma WR
Murchison Falls NP Matiri FR
Kitechura FR
Itwara FR
Table 7 – Relative rankings of sites for Albertine Rift (AR) endemic and globally threatened species
AR endemic species Globally threatened species
High Medium Low
High Virunga NP Rwenzori Mts NP Echuya FR
Itombwe Massif Mt Kabobo Rusizi NR
Kahuzi Biega NP Forest W. of Lake Edward
Kibale NP Mahale Mts NP
Bwindi Impenetrable NP
Nyungwe NP
Medium Kasyoha–Kitomi FR Budongo FR Mafuga FR
Queen Elizabeth NP Kalinzu–Maramagambo Bururi FR
Semuliki NP Lendu Plateau Gombe NP
Kibira NP Mbizi FR
Idjwi Island
Low Mweru-Wantipa NP Bugoma FR Bugungu WR
Murchison Falls NP Semliki WR Kitechura FR
Sumbu NP Kagombe FR Matiri FR
Kyambura WR Itwara FR
Ibambaro FR
Marungu Massif
Karuma WR
190 BIOLOGICAL CONSERVATION 134 (2007) 178–194

Other than this outstanding landscape the following five
areas could be managed on a landscape scale:
1. Nyungwe–Kibira forests: these two existing protected
areas are contiguous across the Rwanda–Burundi border
and it is therefore relatively easy to think about manage-
ment at the landscape scale. Species that may benefit from
management at this scale include chimpanzees, leopards
and golden cats Felis aurata.
2. Murchison Falls National Park – Budongo–Bugoma–
Kagombe–Itwara Forest Reserves – Semliki/Toro Wildlife
Reserve: these sites link Murchison Falls to Semliki Wild-
life Reserve through a corridor of forests reserves, grass-
lands and private forests. This landscape may be
important for gene flow in chimpanzee communities
because few forests in this landscape contain more than
500 individuals. It will require developing corridors that
would link some of the forest reserves, potentially by cre-
ating incentives for private landowners to manage forest
instead of converting it to cultivation.
3. Maiko National Park highlands – Tayna Community
Reserve – Kahuzi Biega National Park – Itombwe Massif:
although not linked by protected areas there is still a fair
amount of natural habitat between these sites and it may
be possible to maintain linkages. It is also important to
maintain the tenuous linkage between the upland and
lowland sectors of Kahuzi Biega National Park. Species
that would benefit from management at this landscape
scale would be elephants and eastern gorillas. There is
currently a programme attempting to create community
reserves that would create the corridors to link these
sites.
4. Mahale Mountains – Katavi–Ugalla: much wild land still
exists to the east of Mahale Mountains National Park and
down towards Katavi National Park. It may be possible to
protect parts of this region to enlarge the park and link
Mahale to other protected areas. Species that would bene-
fit include elephants, chimpanzees, sable Hippotragus niger,
roan Hippotragus equinus and African wild dogs Lycaon
pictus.
5. Marungu Massif and up the western coast of Lake Tang-
anyika to Mount Kabobo. Neither of these sites are pro-
tected and little is known about them. These two sites
probably cannot be linked in a larger landscape but the
area around Marungu still appears to be relatively intact
from satellite imagery (A. Plumptre pers. obs.). A recently
(2005) acquired high resolution satellite image of Mt
Kabobo by WWF shows that significant tracts of montane
forest still exist. Both regions require surveys to assess the
need for protected status and possible conservation of lar-
ger landscapes.
These six landscapes have been identified as core areas by
the regional strategic framework plan that was developed for
the Albertine Rift by conservation NGOs and government pro-
tected area institutions. These landscapes cover most of the
natural habitat in the Albertine Rift and should be thought
of more as political units to help plan for their management,
rather than priorities per se. The priority sites in the Albertine
Rift are those identified in Table 7.
Most assessments of this region have not considered the
lakes within the Albertine Rift. Whether or not they are con-
sidered part of the Albertine Rift, together with Lake Malawi,
they should have greater support from the conservation com-
munity because of their enormous wealth of biodiversity.
Many of the African lakes are poorly known for their fish
fauna and some studies in the early 20th century did not rec-
ognise the diversity of the cichlids at the time they were sur-
veyed (Kaufman et al., 1996; Snoeks, 2000). It is likely many
more species would be identified in Lakes George, Edward
and Albert with further survey work. The number of endemic
species for the Albertine Rift lakes is likely to be more than
the sum of the species endemic to each lake, which already
numbers 366 species.
This analysis focused on species and used species pres-
ence/absence to identify conservation priorities. It is possible
that a focus on subspecies would generate a different result.
We focused on species because there is better agreement on
the taxonomy of species than subspecies and the results are
likely to be more stable. Subspecies tend to be better de-
scribed for mammals and birds than reptiles, amphibians
and plants and it would be complex trying to do a cross taxa
analysis at the subspecies level.
4.2. Conclusions
These results have highlighted the importance of the Alber-
tine Rift within Africa and globally. They have been used to
support the raising of the profile of the Albertine Rift within
African conservation. Our data was used to support Conserva-
tion International’s recent incorporation of the Albertine Rift
within the Eastern Afromontane Hotspot in their re-assess-
ment of global hotspots (Brooks et al., 2004; Plumptre, 2004).
Natural habitat in this region is highly threatened because
of the high density of people (up to 500–600 km
2
) living here
Table 9 – Species richness and numbers of Albertine Rift (AR) endemic and globally threatened species for the Greater
Virunga landscape
Taxon Species richness AR endemic species Threatened species
Mammals 278 30 22
Birds 876 33 17
Reptiles 134 12 1
Amphibians 87 21 10
Fish 81 56 ?
Plants 3552 262 46
Threatened status for freshwater fish species has not been assessed.
BIOLOGICAL CONSERVATION 134 (2007) 178–194 191

(Plumptre and Williamson, 2001). There is a need to focus
more attention on the Albertine Rift to ensure that all the en-
demic and globally threatened species survive. Much of the
Albertine Rift has still been poorly surveyed despite the
wealth of data presented here, and it is probable that many
more species will be discovered if we have the time and re-
sources to invest in biodiversity surveys. Outside protected
areas, natural habitat is being lost at a fairly rapid rate in
the Albertine Rift (Plumptre et al., 2003) and unless we in-
crease the speed and scale of conservation it is likely we will
lose species before they are even discovered.
Acknowledgements
The data compiled and presented here could not have been
completed without a lot of input from experts in the field
and from NGO partners. Royal Museum for Central Africa at
Tervuren, Albertine Rift Conservation Society (ARCOS), Con-
servation International (particularly Tom Butynski), Dian Fos-
sey Gorilla Fund International (particularly Katie Fawcett),
Field Museum at Chicago, Institute for Tropical Forest Conser-
vation (particularly Alastair McNeilage), International Gorilla
Conservation Programme, Makerere University, Missouri
Botanical Gardens (Particularly Roy Gereau), Royal Botanical
Gardens at Kew (particularly Henk Beentje, D. Goyder, and
T. Pearce) Uganda Forest Department (David Hafashimana),
University of Tokyo (Toshisada Nishida), Wildlife Conserva-
tion Society (particularly Isaiah Owiunji, David Nkuutu, Flori-
bert Bujo and Graeme Patterson), and WWF (particularly Neil
Burgess) were amongst the institutions that provided data. In
addition Harald Hinkel, Axel Poulsen, Colin Congdon, Steve
Collins, Alan Gardiner, Neil Baker, Achilles Byaruhanga, Char-
lie Williams, Kim Howell, Simon Stuart, Eberhart Fisher, and
Lauren Chapman all provided information from their own col-
lections/surveys. We are very grateful for all their input and
help. The processes of agreeing on the landscapes presented
here occurred at a workshop organised by ARCOS in February
2003 at which representatives from over 50 protected area
authorities from the Albertine Rift countries, NGOs and indi-
viduals were present and this workshop developed the outline
of the Albertine Rift Strategic Framework document (in press).
The strategic planning process for the Albertine Rift was
funded by the John D. and Catherine T. MacArthur Founda-
tion. Other donors who supported the compilation of these
data included the Daniel K. Thorne Foundation, Wildlife Con-
servation Society, and US Fish and Wildlife Great Ape Conser-
vation Fund. Neil Burgess and Nobby Cordeiro kindly
commented on initial drafts of this manuscript.
REFERENCES
Ackery, P.A., Smith, C.R., Vane-Wright, R.I., 1995. Carcasson’s
African Butterflies – An Annotated Catalogue of the Butterflies
of the Afrotropical Region. The Natural History Museum,
London, UK. p. 803.
Allan, C., 1994. Kyambura game reserve: preliminary results of
Frontier Uganda biological assessment. Uganda Game
Department/Society for Environmental Exploration report.
Anonymous, 1985. Mahale: study for the proposed Mahale
Mountains National park. Final report, JICA, Tokyo.
Ansell, W.F.H., 1978. The Mammals of Zambia. National Parks and
Wildlife Service, Chilanga, Zambia.
Beadle, L.C., 1974. The Inland Waters of Tropical Africa. Longman,
London.
BirdLife International, 2000. Threatened Birds of the World.
Lynx Edicions and BirdLife International, Barcelona/
Cambridge, UK.
Bober, S.O., Herremans, M., Louette, M., Kerbis Peterhans, J.C.,
Bates, J.M., 2001. Geographical and altitudinal distribution of
birds endemic to the Albertine Rift. In: Proc. Pan-African
Ornithological Congress, vol. 10. Ostrich 72, Supplement 1, pp.
189–196.
Brooks, T., Balmford, A., Burgess, N., Fjeldsa, J., Hansen, L.A.,
Moore, J., Rahbek, C., Williams, P., 2001. Towards a
blueprint for conservation in Africa. BioScience 51,
613–624.
Brooks, T., Hoffmann, M., Burgess, N., Plumptre, A., Williams, S.,
Gereau, R.E., Mittermeier, R.A., Stuart, S., 2004. Eastern
afromontane. In: Mittermeier, R.A., Robles-Gil, P., Hoffmann,
M., Pilgrim, J.D., Brooks, T.M., Mittermeier, C.G., Lamoreux, J.L.,
Fonseca, G. (Eds.), Hotspots Revisited: Earth’s Biologically
Richest and Most Endangered Ecoregions, second ed., Cemex,
Mexico, pp. 241–242.
Burgess, N., D’Amico Hales, J., Underwood, E., Dinerstein, E.,
Olson, D., Itoua, I., Schipper, J., Ricketts, T., Newman, K., 2004.
Terrestrial Ecoregions of Africa and Madagascar: a continental
assessment. Island Press, Washington DC. p. 550.
Burtt, B.D., 1934. A botanical reconnaissance in the Virunga
Volcanoes of Kigezi Ruanda. Kivu Bulletin of Miscellaneous
Information 4, 145–165. Royal Botanic Gardens, Kew.
Carcasson, R.H., 1961. The Acraea butterflies of East Africa
(Lepidoptera, Acraeidae). Journal of the East African Natural
History Society (Special Supplement No. 8).
Carcasson, R.H., 1975. The Swallowtail Butterflies of East Africa
(Lepidoptera, Papillionidae). E.W. Classey Ltd., Oxon, UK.
Congdon, T.C.E., Collins, S.C., 1999. Kielland’s Butterflies of
Tanzania. Supplement A.B.R.I. – Lambillionea. Nairobi. p.
143.
Congdon, T.C.E., Gardiner, A., Bampton, I., 2001. Some endemic
butterflies of Kenya, Tanzania and Malawi, ABRI, Nairobi, p. 19.
Cornet d’Elzius, C., 1996. Ecologie, Structure et evolution des
Populations des Grands Mammiferes du Secteur Central du
Parc National des Virunga (Parc National Albert) Zaire (Congo
Belge). 231 pp. Fondation pour Favoriser les Recherches
Scientifiques en Afrique, Belgium.
d’Abrera, B., 1980. Butterflies of the Afrotropical Region.
Lansdowne Press, Melbourne.
d’Abrera, B., 1997. Butterflies of the Afrotropical Region. Part 1,
revised edition. Lansdowne Press, Melbourne.
Davenport, T.R.B., 1996. The Butterflies of Uganda – An Annotated
Checklist. Uganda Forest Department, Kampala, Uganda. p. 48.
Dehn, M., Christiansen, L., 2001. Additions to the known avifauna
of the Rwenzori Mountains National Park in western Uganda.
Scopus 21, 19–22.
Delvingt, W., 1978. Ecologie de l’hippopotame au Parc National
des Virunga. Ph.D. Thesis, Fac. Sc. Agr. Gembloux, Belgium.
Demey, R., Herroelen, P., Pedersen, T., 2000. Additions and
annotations to the avifauna of Congo-Kinshasa (ex Zaire).
Bulletin of the British Ornithological Club 120,
154–172.
de Witte, Z.G.F., 1938. Exploration du Parc National Albert:
mammiferes. Institut des Parcs Nationaux du Congo Belge,
Bruxelles.
de Witte, Z.G.F., 1941. Exploration du Parc National Albert:
batraciens et reptiles. Institut des Parcs Nationaux du Congo
Belge, Bruxelles. 261 pp.
192 BIOLOGICAL CONSERVATION 134 (2007) 178–194

de Witte, Z.G.F., 1965. Les chame
´le
´ons de l’Afrique centrale.
Annales du Musee Royal de l’Afrique Centrale. Sciences
Zoologiques No. 142. Tervuren, Belgium, 200 pp.
Dowsett, R.J., 1990. Survey of the fauna and flora of Nyungwe
Forest, Rwanda. Tauraco Research Report No. 3. 140 pp.
Dowsett, R.J., Dowsett-Lemaire, F., van de Weghe, J.P., undated. Les
oiseaux de la foret de Nyungwe. Checklist published by the
Project Conservation de la foret de Nyungwe.
Drewes, R.C., Vindum, J.V., 1994. Amphibians of the Impenetrable
Forest, southwest Uganda. Journal of African Zoology 108,
55–70.
Drewes, R.C., Vindum, J.V., 1998. Reptiles of the impenetrable
forest (including an addendum and corrigenda to the
amphibian fauna). Unpublished Report, California Academy of
Sciences, San Francisco.
Drewes, R.C., Vindum, J.V., O’Brien, J.P., 1992. Amphibians of the
impenetrable forest, Uganda. Unpublished Report, California
Academy of Sciences, San Francisco.
Eltringham, S.K., 1999. The Hippos. T & D Poyser, Academic Press,
London.
Evans, M., Balmford, A., 1992. Birds in the Ishasha sector of Queen
Elizabeth National Park, Uganda. Scopus 16, 34–49.
Fischer, E., 1996. Die vegetation des Parc National de
Kahuzi-Biega, Sud Kivu, Zaire. Ph.D. thesis, Franz Steiner
Verlag, Stuttgart, 240 pp.
Friedmann, H., Williams, J.G., 1970. The birds of the Kalinzu
Forest, southwestern Ankole, Uganda. Contributions in
science. Series 195 of the Los Angeles County Museum of
Natural History, California.
Gaugris, Y., Prigogine, A., van de Weghe, J.P., 1981. Additions
et corrections a
`l’avifaune du Burundi. Le Gerfaut 71,
3–39.
Gentry, A.H., 1990. Four Neotropical Rainforests. Yale University
Press, New Haven.
Gnoske, T.P., Marks, B.D., 1997. Preliminary report of 1997
avifaunal surveys in SW Uganda: Bwindi Impenetrable
National Park and Echuya Forest Reserve. Unpublished report
to Makerere University and the Field Museum of Chicago.
Groves, C.R., 2003. Drafting a Conservation Blueprint. Island Press,
Washington.
Henning, S.F., 1988. The Charaxinae Butterflies of Africa. Aloe
Books, Johannesburg, South Africa.
Herremans, M., Louette, M., Meirte, D., 2002. The importance of
historical collections for in-situ conservation in Africa.
Abstracts 23°International Ornithological Congress, Beijing.
Hilton-Taylor, C. (compiler), 2000. 2000 IUCN Red List of
Threatened Species. IUCN, Gland, Switzerland. 61 pp.
Hinkel, H., Fischer, E., 1988. Reptiles et amphibians du Rwanda et
leurs environnement. Johannes Gutenberg Universitat, Mainz,
Germany.
Howard, P.C., Davenport, T.R.B., 1996. Biodiversity Reports: vols.
1–33. Uganda Forest Department, Kampala.
Hutterer, R., van der Straeten, E., Verheyen, W.N., 1987. A checklist
of the shrews of Rwanda and biogeographical considerations
of African soricidae. Bonn Zoologishe Beitrage 38, 155–172.
INECN, undated. Les oiseaux du Parc National de la Kibira. INECN,
Bujumbura.
Kalina, J., Butynski, T., 1996. Checklist of the birds of the Bwindi
Impenetrable Forest, Uganda. East African Natural History
Society, Nairobi. 26 pp.
Kaufman, L., Chapman, L.J., Chapman, C.A., 1996. The great lakes.
In: McClanahan, T.R., Young, T.P. (Eds.), East African
Ecosystems and their Conservation. Oxford University Press,
Oxford, pp. 191–215.
Kerbis Peterhans, J.C., 1997. Updated list of small mammals
documented with voucher specimens from Bwindi
Impenetrable National Park and its immediate environs.
Unpublished list, ITFC library.
Kerbis Peterhans, J.C., Austin, P.K., 1996. Small mammal
component of the biological inventory of Kabiranyuma
swamp, Mgahinga National Park, Uganda. Unpublished Report
to ITFC, Uganda.
Kerbis Peterhans, J.C., Kityo, R.M., Stanley, W.T., Austin, P.K., 1996.
Small mammals along an elevational gradient in Rwenzori
Mountains National Park, Uganda. In: Osmaston, H.,
Tukahirwa, J., Basalirwa, C., Nyakaana, J. (Eds.), The Rwenzori
Mountains National Park, Uganda: Exploration, Environment
and Biology. Proceedings of the Rwenzori Conference,
Department of Geography, Makerere University, Uganda.
Kielland, J., 1990. Butterflies of Tanzania. Hill House, Melbourne/
London.
Kingdon, J., 1971–1983. Mammals of East Africa. University of
Chicago Press, Chicago.
Kunkel, I., Kunkel, P., 1969. Contribution a
`la connaissance de
l’avifaune de la foreˆ t du Rugege (Rwanda). Revue de Zoologie
et de Botanique Africaines 79, 327–351.
Larsen, T.B., 1991. The Butterflies of Kenya. Oxford University
Press, Oxford.
Laurent, R.F., 1964. Adaptive modifications in frogs of an isolated
highland fauna in central Africa. Evolution 18, 458–467.
Laurent, R.F., 1972. Exploration du Parc National Albert:
amphibians. Institut des Parcs Nationaux du Congo Belge,
Bruxelles. 125 pp.
Lebrun, J.-P., Stork, A.L., 1991–1997. Enumeration des plantes a
fleurs d’Afrique tropicale. Editions des Conservatoire et jardin
botaniques de la Ville de Geneve, Geneva.
Lock, J.M., 1977. The vegetation of Rwenzori National Park,
Uganda. Botanische Jahrbu
¨cher fu
¨r Systematik.
Pflanzengeschichte und Pflanzengeographie 98,
372–448.
Moreau, R.E., 1943. A contribution to the ornithology of the east
side of Lake Tanganyika. The Ibis 85, 377–412.
Muhlenberg, M., Slowik, J., Steinhauer, B., undated. Parc National
de Kahuzi Biega. Guide book, ICCN/GTZ.
Mwasumbi, L.B., 2000. The diversity of Mbizi Forest Reserve,
Sumbawanga. Unpublished report for the Wildlife
Conservation Society of Tanzania and University of Dar es
Salaam. 54 pp.
Nabanyumya, R.N., 1991. Factors influencing tree species diversity
of the medium altitude tropical moist forests of Kalinzu and
Maramagambo in south western Uganda. Unpublished M.Sc.
Thesis, Makerere University Institute for Environment and
Natural Resources.
Olson, D.M., Dinerstein, E., 1998. The global 200: a representation
approach to conserving the earth’s most biologically valuable
ecoregions. Conservation Biology 12, 502–515.
Omari, I., Hart, J.A., Butynski, T.M., Birhashirwa, N.R., Upoki, A.,
M’Keyo, Y., Bengana, F., Bashonga, M., Bagurubumwe, N., 1999.
The Itombwe Massif, Democratic Republic of Congo: biological
surveys and conservation, with an emphasis on Grauer’s
gorilla and birds endemic to the Albertine Rift. Oryx 33,
301–322.
Patterson, G., Makin, J., 1998. The state of biodiversity in Lake
Tanganyika – A literature review. Natural Resources Institute,
Chatham, UK.
Peace Corps and INECN, undated. The mammals of the Kibira
National Park, Burundi. Checklist of species produced for
ICCN.
Pitman, C., 1938. A Guide to the Snakes of Uganda. Wheldon &
Wesley, Codicote.
Pitman, C.R.S., 1974. A Guide to the Snakes of Uganda, Revised Ed.
Wheldon & Wesley, London. 290 pp.
Plumptre, A.J., 2004. Priority Sites for Conservation in the
Albertine Rift and the Importance of Transboundary
Collaboration to Preserve Landscapes. In: Harmon, D.,
Worboys, G.L. (Eds.), Managing Mountain Protected Areas:
BIOLOGICAL CONSERVATION 134 (2007) 178–194 193

Challenges and Responses for the 21st Century. Andromeda
Editrice, Italy, pp. 233–238.
Plumptre, A.J., Williamson, E.A., 2001. Conservation-oriented
research in the Virunga Region. In: Robbins, M.M., Sicotte, P.,
Stewart, K.J. (Eds.), Mountain Gorillas: Three Decades of
Research at Karisoke. Cambridge University Press, Cambridge,
pp. 361–389.
Plumptre, A.J., Masozera, M., Fashing, P.J., McNeilage, A., Ewango,
C., Kaplin, B.A., Liengola, I., 2002. Biodiversity surveys of the
Nyungwe forest Reserve in SW Rwanda. WCS Working Paper
No. 19, 95 pp.
Plumptre, A.J., Behangana, M., Davenport, T.R.B., Kahindo, C.,
Kityo, R., Ndomba, E., Nkuutu, D., Owiunji, I., Ssegawa, P., Eilu,
G., 2003. The Biodiversity of the Albertine Rift. Albertine Rift
Technical Reports No. 3, p. 105. Available from:
<www.albertinerift.orgnarift-publications>.
Plumptre, A.J., Kujirakwinja, D., Treves, A., Owiunji, I., Rainer, H.,
2007. Transboundary Conservation in the Greater Virunga
Landscape: its importance for Landscape Species. Biological
Conservation 134, 279–287.
Poulsen, A.D., 1997. Plant diversity in forests of western Uganda
and eastern Zaire (preliminary results). AArhus University
reports No. 36. University of Copenhagen, 76 pp.
Prigogine, A., 1953. Contribution a
`l’e
´tude de la faune
ornithologique de la re
´gion a
`l’ouest du lac Edouard. Annales
du Musee Royal de l’Afrique Centrale. Sciences Zoologiques
No. 24. Tervuren, Belgium, 117 pp.
Prigogine, A., 1960. La faune ornithologique du Massif du Mont
Kabobo. Annales du Musee Royal de l’Afrique Centrale.
Sciences Zoologiques No. 85. Tervuren, Belgium, 46 pp.
Prigogine, A., 1967. La faune ornithologique de l’ı
ˆle Idjwi. Revue de
Zoologie et de Botanique Africaines 75, 249–274.
Prigogine, A., 1971–1984. Les oiseaux de l’Itombwe et de son
hinterland. Annales du Musee Royal de l’Afrique
Centrale. Sciences Zoologiques No. 185, 223, 243. Tervuren,
Belgium.
Prigogine, A., 1985. Conservation of the avifauna of the forests of
the Albertine Rift. In: Conservation of Tropical Forest Birds.
International council for Bird Preservation Technical
Publication No. 4. BirdLife International, Cambridge,
pp. 277–295.
Robyns, W., 1948–1955. Flore des Spermatophytes du Parc
National Albert: vols. 1–3. Institut des Parcs Nationaux du
Congo Belge, Bruxelles.
Sanderson, E.W., Redford, K.H., Vedder, A., Coppolillo, P.B., Ward,
S.E., 2002. A conceptual model for conservation planning
based on landscape species requirements. Landscape and
Urban Planning 58, 41–56.
Schiøtz, A., 1999. Treefrogs of Africa. Edition Chimaira, Frankfurt.
Schouteden, H., 1938. Exploration du Parc National Albert:
oiseaux. Institut des Parcs Nationaux du Congo Belge,
Bruxelles. 107 pp.
Schouteden, H., 1949. Contribution a
`l’e
´tude de la faune
ornithologique du Katanga (Congo belge). Review Zoology and
Botany Africa 62, 158–174. and 322–371.
Schouteden, H., 1966. La faune ornithologique du Burundi. Musee
Royal de l’Afrique Centrale. Documentation zoologique No. 11.
Tervuren, Belgium, 81 pp.
Snoeks, J., 2000. How well known is the icthyodiversity of the
large East African Lakes. In: Rossiter, A., Kawanabe, H. (Eds.),
Ancient lakes: Biodiversity Ecology and Evolution, Advances in
Ecological Research, 31. Academic Press, New York, pp. 17–38.
Spawls, S., Howell, K., Drewes, R., Ashe, J., 2002. A field guide to
the reptiles of East Africa. Academic Press, London.
Stanford, C., Msuya, P., 1995. An annotated list of the birds of
Gombe National Park, Tanzania. Scopus 19, 38–46.
Stattersfield, A.J., Crosby, M.J., Long, A.J., Wege, D.C., 1998.
Endemic Bird Areas of the World: priorities for biodiversity
conservationBirdLife International Conservation series No. 7.
BirdLife International, Cambridge.
Stubblefield, L., 1993. Semliki (Toro) Game Reserve: preliminary
results of the Frontier Uganda biological assessment. Uganda
Game Department/Society for Environmental Exploration
report.
Synnott, T.J., 1985. A checklist of the flora of Budongo Forest
Reserve, Uganda, with notes on ecology and phenology. C.F.I.
Occasional Papers No. 27, University of Oxford.
Troupin, G., 1978–1988. Flore du Rwanda: vols. 1–4. Musee Royal
de l’Afrique Centrale. Tervuren and Institut National de
Recherche Scientifique, Butare.
Troupin, G., 1992. Mission d’etude sur la flore de la foret
Nyungwe. Unpublished report to Projet PASP II –
UGZ3 – Nyungwe, Ministere de l’Agriculture de l’Elevage et
des Forets. 70 pp.
Ulfstrand, S., Lamprey, H., 1960. On the birds of the
Kungwe–Mahari area in western Tanganyika territory. Journal
of the East African Natural History Society 23, 223–232.
van der Straeten, E., Kerbis Peterhans, J.C., 1999. Praomys degraaffi,
a new species of Muridae (Mammalia) from central Africa.
South African Journal of Zoology 34, 80–90.
van de Weghe, J.P., Loiselle, B.A., 1987. The bird fauna of Bururi
Forest, Burundi. Le Gerfaut 77, 147–164.
Verheyen, R., 1947. Exploration du Parc National Albert:
oiseaux. Institut des Parcs Nationaux du Congo Belge,
Bruxelles. 87 pp.
Vollesen, K., Bidgood, S., 1996. Kew expedition to Tanzania:
January–May 1994. Unpublished report, 45 pp.
Vollesen, K., Bidgood, S., 1999. Kew expedition to Tanzani:
April–May 1997. Unpublished report, 57 pp.
Vonesh, J.R., 1998. The amphibians and reptiles of Kibale Forest,
Uganda: herpetofaunal survey and ecological study of the
forest floor litter community. Unpublished M.Sc. thesis,
University of Florida.
Weber, W., 1989. Conservation and Development on Zaire–Nile
Divide. An analysis of value conflicts and convergence in the
management of afromontane forests in Rwanda. Ph.D. Thesis,
University of Wisconsin, Madison.
Wilson, R., 1982. The birds of the Parc National des Volcans.
Unpublished checklist, Mountain Gorilla Project, Rwanda.
Wilson, S.E., 1995. Uganda National Parks: Bird and Mammal
Checklists for Ten National Parks in Uganda. National
Biodiversity Databank, Makerere University, Kampala.
Wilson, J.R., Catsis, M.C., 1990. A preliminary survey of the forests
of the Itombwe mountains and the Kahuzi Biega national park
extension, east Zaire: July–September 1989. Unpublished
report to ICCN and WWF.
194 BIOLOGICAL CONSERVATION 134 (2007) 178–194