The biological importance of the Eastern Arc Mountains
of Tanzania and Kenya
*, T.M. Butynski
, N.J. Cordeiro
, N.H. Doggart
, J. Fjeldsa
, K.M. Howell
, S.P. Loader
, J.C. Lovett
, B. Mbilinyi
, M. Menegon
, A. Perkin
, F. Rovero
, W.T. Stanley
, S.N. Stuart
Conservation and Management of the Eastern Arc Mountain Forests (CMEAMF), Forestry and Beekeeping Division, P.O. Box 289,
WWF-USA Conservation Science Programme, 1250 24th St. NW, Washington, DC 20037-1193, USA
Conservation Biology Group, Zoology Department, Cambridge University, CB2 3EJ, UK
Eastern Africa Regional Program, Conservation International, c/o IUCN, P.O. Box 68200, Nairobi 00200, Kenya
Field Museum of Natural History, 1400 S Lake Shore Drive, Chicago, IL 60605, USA
Tanzania Wildlife Research Institute, P.O. Box 661, Arusha, Tanzania
Tanzania Forest Conservation Group, P.O. Box 23410, Dar es Salaam, Tanzania
Zoological Museum, Universitetsparken 15, DK-2100, Copenhagen, Denmark
Zoology Department, University of Dar es Salaam, P.O. Box 35064, Dar es Salaam, Tanzania
Forestry and Beekeeping Division, Ministry of Natural Resources and Tourism, P.O. Box 9372, Dar es Salaam, Tanzania
Department of Zoology, the Natural History Museum, Cromwell Road, SW7 5BD, UK
Environment Department, University of York, Heslington, York YO10 5DD, UK
Sezione di Zoologia dei Vertebrati, Museo Tridentino di Scienze Naturali, Via Calepina 14, I-38100 Trento, Italy
Wildlife Conservation Society, P.O. Box 936, Iringa, Tanzania
Field Museum of Natural History, 1400 S Lake Shore Drive, Chicago, IL 60605, USA
CI/CABS – IUCN/SSC Biodiversity Assessment Initiative, c/o Center for Applied Biodiversity Science, Conservation International, 1919 M
Street NW, Suite 600, Washington, DC 20036, USA
Received 17 June 2005
Received in revised
form 29 June 2006
Accepted 8 August 2006
Available online 12 October 2006
Eastern Arc Mountains
The Eastern Arc Mountains are renown in Africa for high concentrations of endemic
species of animals and plants. Thirteen separate mountain blocks comprise the Eastern
Arc, supporting around 3300 km
of sub-montane, montane and upper montane forest, less
than 30% of the estimated original forested area. At least 96 vertebrate species are endemic,
split as follows: 10 mammal, 19 bird, 29 reptile and 38 amphibian species. This includes
four endemic or nearly endemic species of primate – the Sanje Mangabey, the Iringa Red
Colobus, the Mountain Galago and the new Kipunji monkey that forms its own monotypic
genus. A further 71 vertebrate species are near-endemic. At least 800 vascular plant species
are endemic, almost 10% of these being trees. These endemics include the majority of the
species of African violet – Saintpaulia, a well-known ﬂowering plant in Western households.
An additional 32 species of bryophytes are also endemic. Many hundreds of invertebrates
are also likely to be endemic, with data for butterﬂies, millipedes and dragonﬂies indicating
potential trends in importance. Seventy-one of the endemic or near-endemic vertebrates
are threatened by extinction (8 critical, 27 endangered, 36 vulnerable), with an additional
seven wide ranging threatened species. Hundreds of plant species are also threatened.
0006-3207/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
*Corresponding author: Tel.: +44 1223 440155.
E-mail address: email@example.com (N.D. Burgess).
BIOLOGICAL CONSERVATION 134 (2007) 209–231
available at www.sciencedirect.com
journal homepage: www.elsevier.com/locate/biocon
Most Eastern Arc endemics are closed-forest specialists and comprise taxa with an ancient
history and those of more recent origin, including some possessing ancient afﬁnities with
taxa from West Africa, Madagascar, and even South America and Southeast Asia. Mountain
block prioritisation for biodiversity conservation shows that Udzungwas, East Usambaras
and Ulugurus are the most important blocks, with other important blocks being the Ngurus
and West Usambaras. Rankings are correlated closely with the area of remaining forest.
Most of the remaining forest is found within nearly 150 Government Forest Reserves, with
106 of these managed nationally for water catchment, biodiversity and soil conservation
and where forest exploitation is not allowed. Outside these areas most forest has been
cleared, except in small village burial/sacred sites, a few Village Forest Reserves, and inac-
cessible areas. In most Eastern Arc Mountains the local populations have not encroached
beyond the reserve boundaries to develop farms, but forest resources within the bound-
aries are used for fuel and building materials and some forests are heavily degraded. Fire
is also a problem as it enters and destroys forests during the dry seasons. The future of
the biodiversity on the Eastern Arc Mountains is closely tied to management policies and
capacity of the Tanzania Forestry and Beekeeping Division, Tanzania National Parks
Authority, and Kenya Forest Department. Supporting these agencies in their mandated
job is an essential conservation investment over the longer term.
2006 Elsevier Ltd. All rights reserved.
The phrase ‘Eastern Arc Mountains’, describing a chain of
mountains starting in southern Kenya and progressing
through eastern Tanzania, ﬁrst appeared in print in 1985,
together with a map of the area (Lovett, 1985). In subsequent
years, biological data on the importance of the Eastern Arc
Mountains have been increasingly reﬁned and summarised,
ﬁrst by Lovett and Wasser (1993), and later by Burgess et al.
(1998a,b) and Newmark (2002). These and other data havebeen
used in several global analyses of biodiversity priority to show
that the Eastern Arc Mountains rank among the most impor-
tant areas of the world for the conservation of endemic birds
(ICBP, 1992; Stattersﬁeld et al., 1998), endemic plants (Lovett,
1988; Myers, 1988, 1990; Lovett, 1998a; Mittermeier et al.,
1998; Myers et al., 2000; Lovett et al., 2004), and a combined
set of taxonomic groups (Olson and Dinerstein, 1998; Brooks
et al., 2001; Burgess et al., 2004b). The Eastern Arc is also home
to four endemic or near-endemic species of primates – the
Sanje Mangabey Cercocebus sanjei, Iringa Red Colobus Procolo-
bus gordonorum, the Mountain Galago Galagoides orinus and
the new Kipunji monkey Rungwecebus kipunji that is the sole
representative of its genus (Davenport et al., 2006) – and most
of the known species of African violet – Saintpaulia spp. At-
tempts to factor threat into assessments of conservation prior-
ity across the world and across Africa have also shown that the
Eastern Arc is amongst the most threatened regions of global
biodiversity signiﬁcance and one where the extinction risk to
the fauna and ﬂora is intense, and increasing (Balmford
et al., 2001a,b; Brooks et al., 2002; Burgess et al., 2004a,b).
A number of studies have also looked at patterns and pri-
orities among the 13 separate mountain blocks within the
Eastern Arc (Burgess et al., 1998b; Baker and Baker, 2002; CEPF,
2003), and some have looked at priorities within single blocks
(Rodgers and Homewood, 1982a,b; Johansson et al., 1998;
Dinesen et al., 2000). These studies concluded that some of
the blocks are more important than others, with the East
Usambaras, Ulugurus and Udzungwas consistently being as-
sessed as the most important areas. More recent studies have
shown how the prioritisation can be altered by undertaking
new ﬁeldwork in poorly known blocks, and also that the over-
all ranking of the 13 blocks is correlated with both the forest
area remaining and the degree of study effort (Doggart
et al., 2006).
The primary aim of this paper is to analyse the biological
importance of the Eastern Arc Mountains vertebrate fauna,
as indicated by the number of endemic and near-endemic en-
demic species in the various mountain blocks. We use both
data published in the peer-reviewed literature and that from
unpublished sources provided by taxonomic experts. A sec-
ond aim is to assess the importance of different blocks for
vertebrate species regarded as threatened with extinction
(http://www.redlist.org), with lower signiﬁcance given to this
attribute because the Red List assessment process has not
been completed for reptiles. A third aim is to compile and
present data on other less well known groups to see whether
they share similar patterns of importance. This is done for
trees, and for some invertebrate groups – butterﬂies, milli-
pedes, spiders and dragonﬂies. Prioritisation among the dif-
ferent Eastern Arc Mountain blocks is also examined against
one potential explanatory variable – the area of remaining for-
est. Discussion focuses on hypotheses developed to explain
the exceptional biodiversity of these mountains – which re-
volve around the likely persistence of forest cover over mil-
lions of years and the relative importance of ancient relict
endemics and recently evolved endemics within the Eastern
2. Study area
2.1. Deﬁnition of the Eastern Arc mountains
The Eastern Arc is deﬁned as a chain of ancient crystalline
Precambrian basement mountains, stretching from the Taita
210 BIOLOGICAL CONSERVATION 134 (2007) 209–231
Hills in Kenya to the Udzungwa Mountains in south-central
Tanzania, which were uplifted at least 30 million years ago
and which are under the direct climatic inﬂuence of the
Indian Ocean (Lovett, 1990; Lovett et al., 2004). The Eastern
Arc is also deﬁned to end at the Makambako Gap – a non-
forest region of dry habitat – at the south-western end of
the Udzungwa Mountain range. Forested mountains with a
similar geology are found further south-west of the
Makambako Gap. These ‘Southern Rift’ forests differ from
those of the Eastern Arc because they are not directly under
the Indian Ocean climatic regime and are subject to more
variable convectional rainfall patterns. They also contain
higher percentages of grassland habitat, and while a few
representatives of the ﬂora and fauna are shared with the
Eastern Arc, the ‘Southern Rift’’ forests have much lower
rates of endemism in all taxonomic groups (Lovett, 1993;
Burgess et al., 2004a,b).
Following from the above deﬁnition, there are 13 separate
mountain blocks in the Eastern Arc. These are from north to
south: Taita (including Kasigau; in Kenya), North Pare, South
Pare, West Usambara, East Usambara, Nguu, Nguru, Uluguru,
Ukaguru, Rubeho, Malundwe, Udzungwa, Mahenge (in Tanza-
nia) (Fig. 1;Table 1).
2.2. Forest categorisation
The Eastern Arc Mountains range up to 2635 m in altitude
(Lukwangule Plateau and Kimhandu Peak in the Ulugurus)
and contain a diverse assemblage of habitats. It is expected
that prior to major human inﬂuence on the landscape, the
wetter (eastern and south-eastern) slopes supported a contin-
uous forest cover throughout all elevations, while the drier
(western and north-western) slopes supported deciduous
woodland at lower elevations and evergreen forest only at
higher elevations. Tall evergreen forest was found on the
top plateaus well away from the rain-capturing scarps, as a
consequence of persistent fog over the highlands during the
night. In other parts of the highlands, montane grassland
and heathland dominated. A desiccation-adapted ﬂora
occurred on rocky outcrops.
On the Uluguru Mountains, the forest formations have
been divided into upper montane (1800–2635 m), montane
(1250–1800 m) and sub-montane (800–1250 m) forest zones
´cs, 1976). Elsewhere the same zones are recognised
(Lovett, 1993), but their boundaries occur at somewhat differ-
ent altitudes – depending on inclination of the terrain, rain-
fall, distance from the coast, height of the mountains,
Fig. 1 – Map showing the 13 blocks of the Eastern Arc Mountains of Tanzania and Kenya.
BIOLOGICAL CONSERVATION 134 (2007) 209–231 211
incidence of cloud cover, etc. For example, the forest zone
divisions are at lower elevations in the cloudy and maritime
East Usambara Mountains (Moreau, 1966), whereas they are
much higher in inland rain-shadow areas, where evergreen
forest is limited to top plateaus (>2000 m). At lower elevations
(regarded by Po
´cs, 1976 as below 800 m on the Ulugurus, but
below 500 m elsewhere) the sub-montane forest grades
in species composition and physiognomy into that of the
Table 1 – Location, area (km
) and altitudinal range of forested habitats in the Eastern Arc Mountains of Kenya and
Taita Hills 0325S 3820E 6 3
Not available Not
North Pare 0335–0346S 3733–3740E 151 25
South Pare 0404–0434S 3745–3801E 333 211.1
West Usambara 0420–0507S 3806–3841E 547 220
East Usambara 0445–0520S 3826–3848E 413 450
Nguu 0527–0538S 3736–3732E Included in
Nguru 0527–0613S 3726–3737E 647 328.35
Uluguru 0651–0712S 3736–3745E 528 230
Ukaguru 0619–0635S 3653–3703E 184 155.38
Rubeho 0648–0722S 3634–3658E 499 654
Malundwe Hill 0724S 3718E 6 4.5
Mahenge 0837–0838S 3642–3644E 291 5
Udzungwa 0722–0843S 3507–3658E 2103 1017
Total 5708 3443 3534
a Waiyaki, unpublished (ground survey).
bLovett and Po
´cs (1993) (ground survey).
cCordeiro and Kiure (1995) (ground survey).
dIversen (1991) (aerial photos).
eEvans (1997) (ground survey).
fMoyer (1992) (remote sensing).
˚et al. (1997a,b) (ground survey).
hSvendsen and Hansen (1995) (ground survey).
iJohansson and Sandy (1996) (satellite images and aerial photos).
jSeddon et al. (1995) (ground survey).
kEvans and Anderson (1993) (ground survey).
lLovett and Norton (1989) (ground survey).
mTetlow (1987) (ground survey).
nLovett et al. (2001) (ground survey).
oDoggart et al. (2006) (ground survey).
p Burgess et al. (2002b) (aerial photos and ground survey)
qFrontier-Tanzania (2001–2004) (ground survey).
rLens et al. (2002) (satellite images and ground survey).
sMbilinyi and Kashaigili (2005) (satellite images and ground truthing).
* Not included in the study by Mbilinyi and Kashaigili (2005) and hence the ﬁgure from the previous column was used.
** Including outlying hills such as Mindu, Nguru ya Ndege, Mkungwe, Dindili and Kitulang’halo.
212 BIOLOGICAL CONSERVATION 134 (2007) 209–231
‘transitional’ rainforests. Transitional forests are often
grouped within the lowland Coastal Forests found along the
eastern littoral plain of Africa from Somalia in the north to
Mozambique in the south (White, 1983; Burgess and Clarke,
2000). In reality, no hard boundary exists between these two
forest types (Lovett et al., 2001) and in some mountain blocks
there is a continuum between the Eastern Arc and Coastal
Forest types (e.g. East Usambara, Uluguru, Udzungwa).
2.3. Reserved areas
Two National Parks support Eastern Arc habitats in Tanzania.
The ﬁrst is the Udzungwa Mountains National Park
) which contains large areas of mountain forest
and grassland; the second is Mikumi National Park
) that includes a small area (4 km
) of montane for-
est on Malundwe Hill. Both parks have the internationally
agreed protected area code IUCN II and are managed by
the Tanzania National Parks Authority (TANAPA). Three
other areas of the Eastern Arc are gazetted as Nature Re-
serves, but not given IUCN protected area codes. The ﬁrst
of these is the recently declared (1997) Amani Nature Re-
serve (83 km
) managed by the Forestry and Beekeeping Divi-
sion (FBD) in the East Usambara Mountains, and the second
and third are private Nature Reserves, the ﬁrst within the
Muﬁndi Tea Estate in the Udzungwa Mountains (Lovett and
Moyer, 1992) and the second at Mazumbai in the West
Usambara Mountains (Redhead, 1981).
The majority of the rest of the Eastern Arc forest in Tanza-
nia is found within various different categories of Forest Re-
serve. These are managed under two separate administrative
structures and are not assigned any IUCN protected area
codes. The Tanzania Forestry and Beekeeping Division man-
ages the majority of the larger Forest Reserves for water catch-
ment and biodiversity conservation. These 104 ‘national’
Forest Reserves cover 4718 km
and support a mosaic of forest,
grasslands, and other habitat types (Burgess and Kilahama,
2005). A further 32 Local Authority Forest Reserves (138 km
are managed by the Districts, and approximately six Village
Forest Reserves (25 km
) are managed by villages. In addition
there are seven proposed Forest Reserves (611 km
), and 14
proposed village FRs (22 km
) that are not legally gazetted. Pri-
vately managed forests cover around 177 km
and are found at
Mazumbai (owned by Sokoine University), and within the tea
estates of Ambangulu in the West Usambaras, Muﬁndi in the
Udzungwa Mountains and Amani/Kwamkoro in the East
Usambaras (Tab le 2). Within the human-dominated landscape
outside the Reserves and private estates smaller patches of
forest remain under traditional village authority. Almost every
village has a forest patch for rituals and as a burial grove for its
people, but these are generally under 1 km
in area and the
total area is probably under 100 km
(e.g. Mwihomeke et al.,
3.1. Remaining habitat
The forest cover data presented are derived from Mbilinyi
and Kashaigili (2005), a study undertaken at the Sokoine
Table 2 – Protected areas (National Parks), other reserves (Forest Reserves) and areas of private forest found within the 14
districts of the Eastern Arc Mountains of Tanzania, and number of forestry staff available for management (as of 2004)
Mpwapwa 4 154.6 0 0 0 0 0 5
Kilolo Part of
7 (plus 1
805.5 0 0 0 0 0 66
Muﬁndi 6 218.1 15 15.5 2 2.8 134.5 28
Same 2 197.5 7 (plus 2
74.2 3 (proposed) Unknown 0 27
Mwanga 3 (plus 3
74.1 0 0 0 0 0 14
Kilindi 11 303.4 0 0 0 0 0 12
Lushoto 16 340.1 7 13.6 11 (proposed) 22.1 5 36
Korogwe 8 110.5 0 0 0 0 35.2 26
Muheza 11 (plus 1
NR and 1
315.9 0 0 4 9.9 2.1 25
Kilombero Part of
4 673.4 1 34.7 0 0 0 29
Kilosa Part of
8 801.5 0 0 0 0 0 29
Morogoro 8 356.3 2 0.2 0 0 0 42
Mvomero Part of
9 317.9 0 0 0 0 0 7
Ulanga 7 49.6 0 0 0 0 0 18
Notes: FR = Forest Reserve; NR = Nature Reserve; NP = National Park.
BIOLOGICAL CONSERVATION 134 (2007) 209–231 213
University of Agriculture Remote Sensing and GIS labora-
tory (Morogoro, Tanzania) with technical backing from the
Centre for Applied Biodiversity Science at Conservation
International (Washington, DC, USA) and National Aeronau-
tics and Space Administration (NASA). The methodology to
calculate the forest area in the Eastern Arc is outlined
The satellite images used to assess forest cover were se-
lected from a large number of potential images taken during
the dry season (June–November) and yielding minimum
cloud cover (Table 1). Such images maximise the chances
to resolve differences between evergreen forest habitat and
deciduous (in the dry season) woodland habitat. The best
images were ﬁrst combined into a composite image covering
the Eastern Arc Mountains. Combination involved image rec-
tiﬁcation undertaken using second order polynomial trans-
formation and nearest-neighbourhood interpolation.
Maximum likelihood classiﬁer (MLC) remote sensing classiﬁ-
cation methodologies were then utilised to develop the ini-
tial map of different vegetation types in the Eastern Arc.
The supervised classiﬁcation process involved selecting
training sites on the image which represent known land
classes. Training was an iterative process, whereby the se-
lected training pixels were evaluated by performing an esti-
mated classiﬁcation (ALARM command). These ALARM
results were evaluated and training samples were reﬁned
sequentially. The results of initial ﬁeld veriﬁcation were also
included at this stage. In order to reinforce the visual inter-
pretability of images, a colour composite (Landsat TM bands
4 5 3) was prepared and its contrast was stretched using a
Gaussian distribution function. Furthermore, a 3 ·3 high
pass ﬁlter was applied to the colour composite to further en-
hance linear features, e.g. rivers, and patterns such as culti-
vation. All image processing was carried out using ERDAS
The resulting map was checked for errors. Examples of
typical errors are cloud edges being confused with non-for-
ested areas, water being confused with cloud shadows, etc.
At this stage additional sub-classes were added to the set of
training data or old, problematic sub-classes were modiﬁed.
After this a new MLC was run and the process was repeated
over several iterations until the map had sufﬁciently low er-
ror. Classiﬁed images were then recoded to respective classes
i.e. forest, woodland, plantation, etc.
Veriﬁcation of results was undertaken in two ways, at dif-
ferent times in the process. Firstly, ﬁeld surveys were under-
taken during 2005 in the following Mountain blocks;
Uluguru, Ukaguru, Rubeho, Mahenge, Image and Udzungwa.
This gathered ﬁeld data to help with the initial classiﬁcation
process. Second, draft output maps were sent to nine experts
familiar with different Mountain Blocks and their comments
were used to further reﬁne the classiﬁcation of the habitat
types – especially the differentiation of closed forests and
Data on the vertebrate fauna of the Eastern Arc Mountains are
based on the approach presented in Burgess et al. (1998a,b).In
this scheme, endemic vertebrates are those conﬁned to the
Eastern Arc Mountains as deﬁned here and in the African-
wide assessment of ‘ecoregions’ by the World Wildlife Fund
(USA) (Burgess et al., 2004b, 2006). Near-endemic vertebrates
are found in the Eastern Arc ecoregion (ecoregion 19 of Bur-
gess et al., 2004b), but also occur in one or more of the follow-
ing WWF ecoregions – Northern Inhambane–Zanzibar Coastal
Forest Mosaic (ecoregion 20), Southern Rift Montane Forest–
Grassland Mosaic (ecoregion 74) and/or the East African Mon-
tane Forests (ecoregion 18).
For this paper we have signiﬁcantly updated the biological
data in Burgess et al. (1998a,b) with the assistance of key tax-
onomic specialists. For mammals we added all recently de-
scribed species and new distributional records from
published papers: shrews (Stanley et al., 1998a,b; Stanley
and Hutterer, 2000; Stanley et al., 2005a; Stanley and Olson,
2005), rodents (Stanley et al., 1996, 1998a,b); galagos (Honess
and Bearder, 1996; Bearder et al., 2003; Grubb et al., 2003; Per-
kin et al., 2003), monkeys (Jones et al., 2005); duikers (Rovero
et al., 2005); Rubeho Mountains (Doggart et al., 2006); Mal-
undwe Mountain (Stanley et al., 2005b); Mahenge Mountains
(Frontier-Tanzania, 2001–2004); Uluguru Mountains (Doggart
et al., 2005); Udzungwa Mountains (Frontier Tanzania, 2001);
East Usambara Mountains (Johansson et al., 1998; Frontier
Tanzania, 1999–2002), South Pare Mountains (Stanley et al.,
1996, 1998b). Additional distribution data covering several
large mammals was taken from Cordeiro et al. (2005),Grim-
shaw et al. (1995) and unpublished data from the authors.
For birds we added data for several newly described species
(see Annex 1) and mountain blocks according to the following
sources: Uluguru Mountains (Doggart et al., 2005); Rubeho
Mountains (Doggart et al., 2006); Nguu Mountains (Seddon
et al., 1999a,b); Taita Hills (Brooks et al., 1998; Lens et al.,
2002); Udzungwa Mountains (Fjeldsa
˚, 1999; Butynski and
Ehardt, 2003); Mahenge Mountains (Frontier-Tanzania, 2001–
2004); South Pare Mountains (Baker, 2001); Nguru Mountains
(Romdal, 2001a), West Usambara Mountains (Lovett and
Stuart, 2001); East Usambara Mountains (Cordeiro, 1998;
Seddon et al., 1999a,b). Unpublished data was also provided
by the authors. For amphibians we used the same procedure
of adding recently described species (see Annex 1) and new
distributional data as follows: Uluguru Mountains (Doggart
et al., 2005); Rubeho Mountains (Doggart et al., 2006); Nguu
Mountains (Menegon et al., 2003); Mahenge Mountains (Loader
et al., 2003); Udzungwa Mountains (Poynton, 1998; Poynton
et al., 1998; Menegon and Salvidio, 2005); and unpublished data
from the authors. The reptile data are signiﬁcantly updated
from that in Howell (1993) using Spawls et al. (2001), and more
recently described species (see Annex 1) and new distribu-
tional data as follows: Nguru Mountains; Nguu Mountains
(Menegon et al., 2003); Uluguru Mountains (Doggart et al.,
2005) and unpublished data from the authors. The compiled
data on vertebrate animals in each Eastern Arc mountain
block and nearby forest ecoregions (Annex 1) was used for
various analyses, as described below.
We have excluded from our database vertebrate species
that have an exclusively lowland forest distribution, even if
they are found geographically adjacent to the E
Mountain forests. These species are as follows: one bird (Otus
ireneae), two mammals (Kerivoula africana and Galagoides
cocos), four amphibians (Mertensophryne micranotis;Afrixalus
214 BIOLOGICAL CONSERVATION 134 (2007) 209–231
sylvaticus,Hyperolius rubrovermiculatus and Stephopaedes
usambarae), and two reptiles (Lygodactylus williamsi and Pros-
A list of Eastern Arc endemic large trees and their distribution
was compiled using references cited in Lovett (1998a) supple-
mented by more recent ﬁeld observations by Jon Lovett (An-
nex 2). A large tree was deﬁned as greater than 10 m tall
and/or 20 cm diameter at breast height. The deﬁnition of
near-endemic tree species followed the same methodology
as for vertebrates (see above). Species with disjunct distribu-
tions distant from the Eastern Arc are not included. Species
lists for the Taita Hills, Usambara Mountains, Uluguru Moun-
tains and Udzungwa Mountains are more complete than
other mountain blocks due to greater research activity.
Although not used for analysis we also reviewed information
on endemic bryophyte species (Po
3.4. Analysis of biodiversity priority
Two kinds of ranking methods were used to assess the prior-
ity of the different component blocks of the Eastern Arc
Mountains. The ﬁrst method uses the summed numbers of
strictly endemic and/or near-endemic vertebrates and tree
species for each of the 13 Eastern Arc Mountain blocks. The
second method corrects the species data for the effects of for-
est area and thus aims to reduce the potential bias in the pri-
oritization approach – where larger forests contain more
species simply due to the species–area relationship (Rosen-
Area correction was performed using the equation:
where zis the species–area exponent, Sis the number of spe-
cies, Ais area (km
) and SA is the number of species corrected
for area. We set z= 0.2 as this corresponds to empirical results
for a wide variety of taxa and locations in mainland terrestrial
ecosystems (Rosenzweig, 1995).
3.5. Correlation between biological importance and forest
Across the Eastern Arc Mountains there is great variation in
the area of forest remaining in different mountains. We used
Spearman Rank Correlations to examine whether the biolog-
ical importance of each Eastern Arc Mountain block for ende-
mic vertebrates and trees (uncorrected for area) is explained
by the area of remaining forest habitat.
4.1. Forest area assessment
Our analysis of remote sensing imagery indicates that around
of forest was found in the Eastern Arc Mountains in
the late 1990s/early 2000s. This is a similar estimate to that of
derived by Burgess et al. (1998a,b) using estimates
from a variety of published papers (see Tabl e 1). It is less than
the estimate of 5708 km
for ‘natural forest’ derived from a
landcover map of Tanzania produced using 1995 remote sens-
ing images processed by Hunting Technical Services of the UK
(Newmark, 1998; updated by Newmark, 2002)(Table 1). We use
the estimate of 3534 km
of forest for all further analysis in
this paper. The methods used do not allow us to assess the
quality of that forest, but clearly distinguish natural forest
from woodland and tree plantations.
4.2. Eastern Arc endemic species
Ninety-six species of vertebrates are endemic to the Eastern
Arc Mountains (Annex 1). Of these, 73 (76%) are dense forest
specialists and only two (2%) occupy non-forest habitats.
Most of the endemic species are sedentary within the Eastern
Arc forests, with only a few of the bird species undertaking
seasonal migrations as temperatures fall in these mountains
during the austral winter months (July–September) (Burgess
and Mlingwa, 2000; Romdal, 2001b).
A further 71 species of vertebrate are near-endemic to the
Eastern Arc Mountains. Of these, 28 are also found in the
lowland Coastal Forests, 46 also range south into the South-
ern Rift, and 10 are also found on the younger volcanic high-
lands of northern Tanzania and/or Kenya (some occur in
more than one of these three broader regions). The majority
of these species are also forest-dependent (45 species; 63%),
but some (6 species; 8.5%) occupy non-forest habitats. A few
of the near-endemic species also migrate out of the Eastern
Arc Mountains, for example the Blue Swallow Hirundo
A total of 68 endemic or near-endemic trees are recorded for
the Eastern Arc. These tree species occur throughout the ele-
vation and moisture range of the forests, but higher numbers
are found in wetter forests. Many of the lowland forest spe-
cies of the Eastern Arc also occur in coastal forests to the east
4.3. Threatened species in the Eastern Arc mountains
Most of the birds, mammals and amphibians of the Eastern
Arc have been assessed against the IUCN Red List criteria –
quantifying their degree of threat (http://www.redlist.org,
November 2005). Seventy-one of the Eastern Arc endemic or
near-endemic vertebrates are threatened by extinction (8 crit-
ical, 27 endangered, 36 vulnerable (and 8 data deﬁcient and
two near-threatened)) with seven other more wide ranging
threatened species. Some of the newly described species (par-
ticularly of birds and mammals) are not yet included on the
Red List because they have not been assessed. These omis-
sions include three newly described species of mammal (Con-
gosorex phillipsorum,Hylomyscus arcimontensis and R. kipunji,
the latter being an unusual new monotypic genus: Davenport
et al., 2006), species of Nectarinia sunbirds (Bowie et al., 2004)
and Turdus thrushes (Bowie et al., 2005), and an akalat – Shep-
pardia aurantiithorax (Beresford et al., 2004). Most reptiles have
also not been assessed against Red List criteria and many
BIOLOGICAL CONSERVATION 134 (2007) 209–231 215
would be expected to qualify given their small ranges and the
extensive collection of some (e.g. chameleons) for the pet
The Red List of threatened tree species for the Eastern Arc
Mountains was reviewed in 2003 and was regarded as too
incomplete to be used for analysis (CEPF, 2003; Gereau and
Luke, 2003). Due to this shortcoming we did not attempt an
analysis of threatened tree species among the various Eastern
Arc Mountain blocks. However, we know that many of the
Eastern Arc endemic tree species are threatened and some
may have already become extinct – for example the tree Platy-
pterocarpus tanganyikensis (a monotypic genus) was only
known from an area of forest in the West Usambara Moun-
tains that was cleared shortly after Tanzania’s independence
in 1963. This tree species has not been relocated since (Lovett
and Stuart, 2001).
4.4. Biodiversity priorities
The Udzungwa Mountains have the largest number of single-
block endemic (found in no other forest blocks) vertebrate
species (17 species), followed by the Uluguru (13 species),
the Taita Hills (6 species), the West Usambara (5 species)
and the East Usambara (4 species) (Fig. 2a). All other blocks
have two or less single block endemic species, and four blocks
have none at all. When corrected for area, the Uluguru Moun-
tains and the Taita Hills rise marginally above the Udzungwas
in terms of single-block endemic vertebrates per unit area of
forest habitat remaining (Fig. 2a).
In terms of Eastern Arc endemic vertebrate species, the
Uluguru Mountains have the largest number of vertebrate
species (44 species), followed by the Udzungwa (41 species),
the East Usambara (35 species), the West Usambara (22 spe-
cies), the Nguru (20 species) and the Rubeho (12 species)
(Fig. 2b). All other sites have 10 or fewer Eastern Arc endemic
Number of single block endemics
Single block endemics per sq km
Numberof endemic species
Numberof near-endemic species
Endemic species per sq km
Number of endemic and near-
Endemic and near-endemic
species per sq km
Near-endemic species per sq km
Number of threatened species
Threatened species per sq km
Numberof endemic and near-endemic
Endemic and near-endemictrees per sq
Fig. 2 – Ranked importance for vertebrates and large trees of the 13 mountain blocks of the Eastern Arc Mountains (grey
bars = uncorrected species data; black lines and diamonds = species data corrected for forest area). (a) Single block endemic
vertebrate species; (b) Eastern Arc endemic vertebrate species; (c) Endemic and near-endemic vertebrate species; (d) Near-
endemic vertebrate species; (e) Threatened vertebrate species; (f) Numbers of endemic and near-endemic tree species.
Mountain blocks are arranged left to right from north to south in the Eastern Arc.
216 BIOLOGICAL CONSERVATION 134 (2007) 209–231
vertebrate species, with the North and South Pare Mountains
and Mahenge Mountains being particularly impoverished.
When corrected for variations in forest area, the Uluguru
Mountains become of the highest importance for the conser-
vation of Eastern Arc endemic vertebrate species (Fig. 2b).
When endemic and near-endemic vertebrates are com-
bined, the Udzungwa Mountains have the largest number of
vertebrate species (96 species), followed by the Uluguru (81
species), the East Usambara (77 species), the Nguru (52 spe-
cies), the West Usambara(48 species), the Rubeho (35 species),
the Ukaguru and Nguu (27 species) (Fig. 2c). Of lowest impor-
tance on this scale is Malundwe Mountain with two Eastern
Arc endemic or near-endemic species (but virtually no
research has been done there). When adjusted for differences
in forest area, the Uluguru and East Usambara Mountains are
the most important sites for endemic and near-endemic spe-
cies (Fig. 2c). Similar patterns are seen with near-endemic
species (Fig. 2d).
Finally, when the number of Threatened species is used as
the measure of importance, the following priorities are
obtained: Most important is the Udzungwa Mountains (40
species), followed by the Uluguru (29 species) and East Usam-
bara (28 species), and West Usambara (21 species) (Fig. 2e).
When adjusted for variations in forest area, the order of
importance of these mountains remains the same (Fig. 2e).
The number of endemic and near endemic trees varies con-
siderably among blocks across the Eastern Arc Mountains
(Fig. 2f). The highest number of Eastern Arc endemic/near
endemic trees are found in the East Usambara (40 species),
Udzungwa (37 species), West Usambara (27 species) and
Uluguru (26 species) Mountains (Fig. 2f; Annex 2), which
are all well studied blocks. This ranking changes a little
when the number of endemic and near-endemic trees is cor-
rected for the remaining area of forest in these blocks
(Fig. 2f). Under this analysis the most important forests are
those of the East Usambara Mountains, followed by the West
Usambara, the Udzungwa, the Uluguru and the Nguru
Mountains. The small forests of the Taita Hills are also
important in terms of numbers of species in a very small
area of forest, as are those of Mahenge. The other mountain
blocks are all poorly known, and none are known to contain
more than six endemic or near-endemic trees, with Rubeho
and North Pare having no known endemics. It is likely that
the number of endemic trees in the Nguu, Ukaguru, Rubeho,
North Pare and Mahenge Mountains will increase as botani-
cal exploration continues.
4.5. Relationship between vertebrate species importance
and forest area
There is a high correlation between the vertebrate importance
of an Eastern Arc Mountain block and the area of forest
remaining. Positive correlations are found between forest
area and (1) numbers of endemic plus near-endemic species
(Spearman rank correlation coefﬁcient r
= 0.862; P< 0.0001),
(2) numbers of Eastern Arc endemic species (Spearman
= 0.831; P< 0.0001), and (3) numbers of Red List species
= 0.815; P< 0.001). The Red List correlation is
easily explained as a small geographical range is one of the
criteria within the red listing process.
This paper demonstrates that the Eastern Arc Mountains are
of exceptional importance for endemic species of vertebrates
and trees in Africa. It also shows how the mountain blocks
within the Eastern Arc attain different importance scores
for the conservation of species vertebrates and trees. The
Uluguru Mountains, East Usambara Mountains and Udzung-
wa Mountains consistently emerge as the most important
blocks for vertebrates and trees, whereas the Nguru and West
Usambara Mountains also ﬁgure as important for tree spe-
cies. The prioritisation results are correlated strongly with
the area of remaining forest.
In the following sections we discuss whether (1) the high
rates of endemism seen in vertebrates and trees are also
found in other taxonomic groups, (2) outline some of the
hypotheses put forth to explain why there is so much ende-
mism in the Eastern Arc, and (3) present some suggestions
on where survey effort needs to be focussed in the future to
obtain a better picture of the patterns of endemism in the
Eastern Arc. Finally, we present the conservation challenges
facing the Eastern Arc Mountains of Tanzania and the work
that is ongoing to try and ensure the long term conservation
of the natural habitats of the Eastern Arc – and hence the sur-
vival of the endemic species that rely on them.
5.1. Additional endemic vertebrates and plants
We still do not have a complete inventory of the endemic spe-
cies of vertebrates and plants in the Eastern Arc Mountains.
New species (and genera) are still regularly being discovered.
Within the plants this includes new genera like Kihansia
(Cheek, 2004), and new species in well-known groups such
as coffee (Davis and Mvungi, 2004). More than 15 new verte-
brate species are also in the process of scientiﬁc description,
including: one Bradypodion and one Urocotyledon from Ma-
henge, two new Nectophrynoides and one Probreviceps from
Ukaguru (one of the Nectophrynoides from Ukaguru has re-
cently been named as N. laticeps –Channing et al., 2005), three
new Nectophrynoides from Uluguru, two Nectophrynoides, three
Callulina, one Arthroleptis and one Probreviceps from Nguru,
and one Afrixalus and one Arthroleptis from Udzungwa.
Detailed DNA and morphological analysis of existing spe-
cies also seems likely to further increase the number of taxa
within these mountains. As examples, molecular (and mor-
phological) variation in bird species distributed across a num-
ber of mountain blocks indicates that they should be split into
allo- or parapatric component species; namely Artisornis met-
opias,Batis mixta (with populations from Ukaguru and Ulug-
uru towards the south west representing an unnamed
species), Modulatrix stictigula,Illadopsis ruﬁpennis and Stactola-
ema olivacea. And ﬁnally in the mammals, there is a distinctive
sub-species of the genet Genetta servalina lowei that is conﬁned
to the Udzungwa Mountains (De Luca and Mpunga, 2002),
Kanga Forest Reserve in the Nguru Mountains and Uluguru
North in the Uluguru Mountains (F. Rovero, unpublished). This
may also, with further work, turn out to be a full species.
BIOLOGICAL CONSERVATION 134 (2007) 209–231 217
5.2. Endemism in other taxonomic groups
The invertebrate fauna of the Eastern Arc is more poorly
known than the vertebrate fauna. However, the available
information suggests that many species of invertebrate are
conﬁned to a single Eastern Arc Mountain block. For example,
Scharff (1992) shows that single site endemism for linyphiid
spiders is over 80%. Moreover, for carabid beetles the Uluguru
Mountains have 95% endemism (Basilewsky, 1962, 1976), and
for harvestmen arachnids this site has 88% endemism
(Scharff et al., 1981). Some of the patterns known for individ-
ual invertebrate groups are outlined below.
5.2.2. Odonata (dragonﬂies and damselﬂies)
Three odonate species are endemic to the Eastern Arc (Platycy-
pha auripes,Amanipodagrion gilliesi and Micromacromia miraculosa
– the last two are East Usambara endemics). Two near-endemic
species found in the Eastern Arc are Umma declivium (Eastern
Arc and north Malawi) and Chlorocnemis abbotti (Eastern Arc
and Kilimanjaro) (Clausnitzer, 2001). Some of these represent
genera that are more widespread in the Central and West Afri-
can forests. The endemic Eastern Arc Odonata species are
found in forest habitats and breed in montane streams, or in
small waterﬁlled holes in tree-trunks. Three coastal forest
endemics that may perhaps range into the lowlands of the East-
ern Arc are Coryphagrion grandis (Gondwana relict with nearest
relatives in Central and South America), Hadrothemis scabrifrons
(relict form also found in coastal Gabon and Cameroon), and
Thermochoria jeanneli (coastal swamp forest) (Clausnitzer, 2001).
5.2.3. Lepidoptera (Butterﬂies and moths)
At least 43 species of butterﬂies are endemic to the Eastern
Arc and contiguous forests in their foothills (De Jong and
Congdon, 1993; Congdon et al., 2001). A further 35 species
are only found on the higher altitude grasslands of the East-
ern Arc and further south into the Southern Highlands of Tan-
zania and into Malawi. The most important Eastern Arc
blocks in terms of endemic butterﬂies are the Rubeho (13 spe-
cies), Udzungwa (9 species), Usambara (7 species), Uluguru (7
species), and Nguru (4 species). The forest butterﬂy fauna also
has genera that are representative of groups which are more
diverse in the Central and West African rainforests.
Hoffman (1993) outlined the state of knowledge of the Eastern
Arc millipede fauna, showing that the East Usambaras,
Udzungwas and Ulugurus (the only areas where inventories
have been compiled) support at least 26 species and 10 genera
endemic to one or other of these mountains. New collections
from the East Usambara Mountains (Frontier Tanzania, 1999–
2002), Uluguru Mountains (Doggart et al., 2005) and Udzung-
wa Mountains (Frontier Tanzania, 2001) hold additional new
genera and species (Hoffman, pers. comm.). It is likely that
the number of endemic genera and species will rise signiﬁ-
cantly with further research.
The Eastern Arc Mountains support a diverse assemblage of
bryophytes, with around 700 species recorded (Po
At least 32 species are endemic (5%). Although this level
of endemism is low compared with vascular plants, it is
high compared with the bryophyte ﬂora of many other
areas. A number of monotypic endemic genera are also
present, for example Cladolejeunea and Neorutenbergia. A
notable feature of the bryoﬂora is the high number (45 spe-
cies, 6%) of Lemurian (Madagascan) species within the
assemblage, which reaches its peak in the Uluguru Moun-
tains (40 species). The bryoﬂora of the Usambara and Ulug-
uru Mountains is quite well known, but information is
scanty to non-existent for the other Eastern Arc mountain
5.3. Possible reasons for the exceptional rates of endemism
Why is there such a concentration of endemism in the East-
ern Arc Mountains? High level of endemism in East Africa
has traditionally been explained by ancient, isolating pro-
cesses related to the uplift of mountains during the Miocene
(Lovett, 1993). These isolating processes would be predicted
to result in species replacement patterns that are congruent
across many evolutionary lineages, something that does not
seem to be supported by recent studies (Bowie, 2003; Bowie
et al., 2004). A ﬁrst step towards understanding the processes
responsible for such high endemism in the Eastern Arc has
been taken by Jetz et al. (2004), who examined whether ende-
mism in an area exceeds what can be explained from simple
null models based on stochastic effects. Jetz et al. (2004) show
that the Eastern Arc Mountains, and the Usambara and Ud-
zungwa Mountains in particular, have signiﬁcantly more en-
demic species than expected by chance, or by environment
and topography alone. These results were interpreted as a
consequence of stable local conditions, but the analysis could
not separate to what degree the pattern can be attributed due
to local speciation or to species persistence (locally low
extinction or high speciation rates). Other kinds of evidence
provide information about the probable historical processes
operating in the Eastern Arc.
5.3.1. The Arc is as a centre of speciation
One of the possible explanations for the high number of
endemics in the Eastern Arc is that the area has had elevated
rates of speciation in situ. Various plant groups show remark-
able local radiations, for instance Impatiens spp. in the Ulug-
uru and Begonia spp. in the Udzungwa Mountains. For
Saintpaulia there is a strong support for recent diversiﬁcation
of local populations in the Usambara Mountains (Lindqvist
and Albert, 2001). Other on-going studies suggest a strong de-
gree of recent genetic differentiation of populations of verte-
brates inhabiting different highlands (e.g. Bowie, 2003 and in
litt. for birds; Perkin et al., 2003 for galagos; Matthee et al.,
2004 for leaf chameleons; Loader et al., 2004 in litt. for
amphibians; E. Verheyen in litt. for rodents). Molecular stud-
ies also suggest both a considerable amount of gene-ﬂow be-
tween populations inhabiting different tracts of highland
forest, but also some gene ﬂow breaks, which in some cases
appear to correspond to physical barriers (Beresford et al.,
2004; Stanley and Olson, 2005), but in other cases are situated
in an area with no obvious physical barriers (Bowie et al.,
˚et al., in press).
218 BIOLOGICAL CONSERVATION 134 (2007) 209–231
5.3.2. The Eastern Arc is as an area where species avoid
It has been suggested that the forest habitats have persisted in
the EAM for a prolonged period, even during particularly ex-
treme climatic phases (Lovett, 1993). This ecological stability is
thought to have promoted the persistence forest species and
overall reducing rates of extinction (Lovett and Wasser, 1993;
˚and Lovett, 1997). The direct evidence for permanence
of forest is lacking, but there is strong circumstantial evidence.
Firstly, a number of the genera and species in the Eastern
Arc Mountains are known to be genetically ancient. For exam-
ple, DNA analysis of forest birds indicates that some species
derive from lineages stretching back to the early Miocene
(c.f. 25 million years ago) (Fjeldsa
˚, 1994; Fjeldsa
1997; Barker et al., 2004; Fuchs et al., 2005). Some of these spe-
cies have their strongest afﬁnities with others in forested
areas in Southeast Asia; for example the Udzungwa partridge
Xenoperdix udzungwensis (Dinesen et al., 1994). Others may
date back to when there was forest cover across Africa, in
the middle and upper Miocene (Axelrod and Raven, 1978)as
DNA analyses of some of the isolated eastern bird subspecies
show they are divergent from their relatives in the Central
Congolian forests (Roy, 1997; Roy et al., 1997; Beresford, 2003).
Secondly, a number of taxonomic groups found in the EAM
contain members (or sometimes the entire group) that are of-
ten referred to as being ‘primitive’, i.e. extant members are
found on more basally branching parts of a phylogenetic tree.
For example, in mammals, the elephant shrews and galagos
are often considered, respectively, as early diverging macros-
celidean and prosimian lineages (Kingdon and Howell, 2005;
Bayes, 1998; Martin, 2003; Fjeldsa
˚et al., 2005; Masters et al.,
2005). These ‘primitive’ groups that are also referred to as an-
cient relict lineages, are well represented in the Eastern Arc
forests and, therefore, suggest a prolonged history of forest
habitats (Burgess et al., 1998a,b). Similarly, the amphibian fau-
na of the Eastern Arc contains a diverse number of taxa, many
of which are thought to share afﬁnities with species found re-
stricted to distant montane regions of Africa (e.g. bufonids,
brevicipitines and caecilan lineages) (Nussbaum, 1985; Poyn-
ton, 1999; Wilkinson et al., 2002). Within the invertebrates,
Hochkirch (1998) presents information that indicates an an-
cient history for the grasshoppers of the Eastern Arc. In addi-
tion the mollusc fauna contains evidence of ancient faunal
connections to Madagascar (Emberton et al., 1997), as does
the vascular plant ﬂora (Lovett and Friis, 1996) and the bryo-
phyte ﬂora (Po
´cs, 1998). Further evidence of the ancient nature
of the Eastern Arc fauna is provided by comparisons with the
visually similar forests at comparable altitudes on the geolog-
ically recent (less than 2 million years old) volcanoes of the re-
gion (e.g. Kilimanjaro, Meru, Hanang). All the Eastern Arc
forested blocks have much higher rates of endemism than
those of these volcanic mountains (see e.g. Scharff, 1992; Po
1998) – and most importantly - the relict species and biogeo-
graphical connections to Asia, Congo Basin and Madagascar
are lacking in the ﬂora and fauna of the recent volcanoes.
Thirdly, despite there being strong evidence that climatic re-
gimes in East Africa highly ﬂuctuated, and as a result so did the
ﬂoral composition (Trauth et al., 2005), it is likely that montane
habitats, such as those in EAM, were subject to less intensive
climatic changes and remained ecologically stable. The
elevated topography afforded montane regions more stable
climates (Lovett, 1993), buffering forest habitats during extreme
arid phases. Marine drill-core data has suggested that the
coastal waters of Tanzania were less inﬂuenced by Pleistocene
climatic changes than elsewhere (Prell et al., 1980), maintaining
warm and seasonally humid conditions in the coastal regions
and presumably the near-shore Eastern Arc Mountains. Long-
term forest stability, as indicated by fossils or climatic indica-
tors has not been investigated directly from any site in the
Eastern Arc, and this remains an important area of future re-
search for understanding the persistence of forest habitats in
the area. Within the period covered by rainfall records, the
African climate has shown marked annual ﬂuctuations in both
wet and dry areas, with the climate in some places having a
much greater inter-annual variability than others (Nicholson,
1994; Owen et al., 1990; Trauth et al., 2005). However, the East-
ern Arc Mountains stand out for having a stable recent climate
within an analysis of 10 years meteorological satellite climate
data, and it has been postulated that this stability has persisted
over the long term (Fjeldsa
˚et al., 1997a,b).
5.4. Conservation priorities
Among the 13 Eastern Arc Mountain blocks the most impor-
tant blocks seem robust to the addition of new data. The top
three blocks have not changed since the analysis of Burgess
et al. (1998a,b) – and do not change when using alternative for-
est area estimates (e.g. Newmark, 1998, 2002) – with the East
Usambara, Uluguru and Udzungwa consistently being the
most important blocks. Plants also indicate the same three
areas as having the highest importance; for example the Ulug-
uru has more than 135 endemic plant species (Burgess et al.,
2002) and the East Usambaras more than 100 endemic species
(Iversen, 1991). The ranking of Eastern Arc blocks further down
the prioritized list has, however, been inﬂuenced by the dis-
covery of new species through undertaking new research in
poorly known mountain areas – e.g. the Rubeho Mountains
(Doggart et al., 2006). This new research has resulted in some
areas (notably the Nguru and the Rubeho Mountains) becom-
ing more important for conservation investment.
5.5. Conservation issues
The majority of the Eastern Arc Mountains are found in Tan-
zania and a number of issues affect their effective conserva-
tion. These issues are similar to those affecting the Taita
Hills of Kenya, where only 3 km
of fragmented forest re-
mains (Lens et al., 2002).
In Tanzania, the future of the biodiversity of the Eastern
Arc Mountains is closely tied to the management capability
and approach of the Forestry and Beekeeping Division (FBD)
of the Ministry of Natural Resources and Tourism, the (parast-
atal) Tanzanian National Parks Authority (TANAPA), and the
District Natural Resource Departments. The majority of the
remaining forest is managed by these agencies. Ultimately,
the conservation of the forests will also depend on the atti-
tudes, wishes and practices of the hundreds of rural commu-
nities living adjacent to the reserves.
BIOLOGICAL CONSERVATION 134 (2007) 209–231 219
The Forestry and Beekeeping Division of the Tanzanian
government is working to improve the conservation status
of the Eastern Arc Mountains in a number of ways. Starting
from 2004, the Tanzanian government has initiated the
process to include the Eastern Arc Mountains on the list
of natural World Heritage Sites. The initial proposal was
submitted in 2005 and a letter received from UNESCO in
January 2006 conﬁrmed that the Eastern Arc has been
placed on the Tentative List of World Heritage Sites. The
government is now working towards developing and sub-
mitting the full proposal dossier for this site – which will
be made up on a number of different forest areas across
In addition Forestry and Beekeeping Division has also
undertaken a detailed analysis of the biological values of
the various blocks and reserves within the Eastern Arc
and has identiﬁed a number of areas that might be up-
graded from ‘protective Forest Reserves’ to ‘nature Forest
Reserves’. These are the Nilo Forest Reserve in the East
Usambara Mountains, the Uluguru North and Uluguru
South Forest Reserves in the Uluguru Mountains, the West
Kilombero Scarp Forest Reserve in the Udzungwa Moun-
tains, and the Nguru South Forest Reserve in the Nguru
Mountains. Active attempts to upgrade the status of these
areas to Nature Reserve are now underway for all but the
South Nguru FR.
To further recognise the importance of the network of For-
est Reserves in the Eastern Arc, the Forestry and Beekeeping
Division is working with the IUCN World Commission on Pro-
tected Areas (WCPA) to code each Forest Reserve in the East-
ern Arc according to IUCN protected area categories. Over 100
reserves have now been provisionally coded against these
categories, with most falling under category IV (habitat/spe-
cies management area). The full list and an accompanying
map will be submitted to IUCN WCPA during 2006.
Finally, the government is also working to gazette some of
the reserves that have been proposed for a number of years,
but never legally declared. There are seven proposed reserves
in the North Pare, South Pare, East Usambara and Udzungwa
Mountains that cover around 62,000 ha of Eastern Arc forest,
grassland and some woodland habitats. Their declaration
would be an important addition to the network of reserves
in this area (http://www.easternarc.or.tz).
A government analysis of problems affecting the manage-
ment of the Eastern Arc reserve network in Tanzania has
shown that the key constraints to effective conservation are
the lack of long-term availability of funding and inadequate
management capacity. Apart from meagre salary support,
the Tanzanian government (both central and local) provides
about $50,000 USD as operational funding to the 340 govern-
ment staff who are tasked with the management of over
of Forest Reserves of the Eastern Arc (Burgess and
Kilahama, 2005) (Tab l e 2 ). The funds available from TANAPA
to its two National Parks containing Eastern Arc mountain
forest (Udzungwa and a small area in Mikumi) are much
greater than that allocated by FBD.
We believe that several possibilities exist to expand sus-
tainable ﬁnancing of Eastern Arc Mountains conservation
within Tanzania and Kenya. One promising avenue involves
payments for ecosystems services, which would create
mechanisms that return funds from water users to the for-
est managers and surrounding communities. The Eastern
Arc provides drinking water for at least 60% of the urban
population of Tanzania, and is the source for over 90% of
the nation’s hydroelectricity generation capacity (which is
around 50% of total power production). A recent calculation
estimates the annual total economic value of the Eastern
Arc as at least $620 million (Government of Tanzania,
2004), but the water values are not well reﬂected in these
estimations and the actual values to the Tanzanian econ-
omy might be signiﬁcantly greater. Another funding avenue
that is developing slowly is to direct the tourism away from
the northern circuit of national parks like the Serengeti,
Ngorongoro Crater and Tarangire systems toward the East-
ern Arc Mountains and coastal zone. This venture requires
adequate global and national marketing, where culture and
biodiversity other than large mammals offer tourists a dif-
ferent type of exploration that could generate money for lo-
cal economies. The government and private sectors are
working on this aspect of tourism for the Eastern Arc
Mountains, and as more local communities are involved,
the more likely the harmonisation of conservation will be
In conclusion, the Eastern Arc Mountains are an area of
exceptional importance for biodiversity conservation. The
majority of the forest is located within national Forest Re-
serves that are managed for water catchment. Despite the
pressures, Forest Reserve boundaries, where they are clearly
deﬁned, are mostly respected by the local people. However,
the funds available to the managers of the Forest Reserves
are insufﬁcient. This makes management problematic and
allows signiﬁcant illegal activities to take place within the
forests (timber cutting etc). The future of the forest re-
source on the Eastern Arc Mountains is tied to the manage-
ment capacity of the Forest and Beekeeping Division
(catchment forest section), to the District Natural Resource
ofﬁces in the 14 Districts across the Eastern Arc, and to
the attitudes and practices of the hundreds of thousands
of people who live within a few kilometres of the forest
Louis A. Hansen in Denmark is thanked for his input to
checking Table 1 of this paper and providing freely his knowl-
edge of the area. We thank all the researchers who have
worked in the Eastern Arc Mountain forests. The longest term
efforts to understand the values of these forests have come
from the University of Dar es Salaam in Tanzania, the Univer-
sity of Copenhagen in Denmark, York University in the UK
and the Frontier-Tanzania expedition programmes of the
Society for Environmental Exploration. We also thank the
organisers of the Society for Conservation Biology meeting
in New York 2004 for the opportunity to present this paper. Fi-
nally, we thank the managers of the Forest Reserves and Na-
tional Parks in the Eastern Arc Mountains as without their
efforts the biodiversity values in these mountains would be
very signiﬁcantly less than is found today. Two anonymous
reviewers and Tim Caro greatly improved the quality of this
220 BIOLOGICAL CONSERVATION 134 (2007) 209–231
Appendix 1. Distribution of endemic and near-endemic vertebrates across the Eastern Arc Mountains
Species Authority Ende-
Forest Taita North
Nguu Nguru Uluguru Ukaguru Rubeho Malundwe Mahenge Udzungwa Coastal
Andropadus fusciceps Shelley (1893)
NE LC FF x x x x x x x x
Andropadus masukuensis Shelley (1897)
NE LC FF x x x x x x x x x x x
Andropadus milanjensis Shelley (1896)
NE LC FF x x x x x x x x x x x x x
Anthreptes pallidigaster Reichenow (1905) NE EN F x xx
Anthreptes rubritorques Sclater and Moreau (1935) E VU FF x x x x x x
Apalis chariessa Reichenow (1879) NE VU FF xxxx
Apalis chapini Friedmann (1928) NE LC FF x x x x x x
Apalis fuscigularis Moreau (1938) E CR FF x
Arcanator orostruthus Vincent (1933) NE LC FF x xx
Artisornis moreaui Sclater (1931) NE CR FF x x
Artisornis metopias Reichenow (1907) NE LC FF x x x x x x x x
Batis mixta Shelley (1889) NE LC FF x x x x x x xn xn xn xn x x xn
Bubo vosseleri Reichenow (1907) E VU FF xu x x x x x x
Cisticola njombe Lynes (1933) NE LC NF xx x
Cisticola nigriloris Shelley (1897) NE LC NF xx x
Cinnyricinclus femoralis Richmond (1897) NE VU FF x x x
Hyliota usambarae Sclater (1932) E EN F x x
Laniarius fuelleborni Reichenow (1900) NE LC FF x x x x x x x
Lanius marwitzi Reichenow (1901) NE LC f xx x x
Malaconotus alius Friedmann (1927) E EN FF x
Modulatrix stictigula Reichenow (1906) NE LC FF x x x x x x x x
Nectarinia loveridgei Hartert (1922) E LC FF x
Nectarinia moreaui Sclater (1933) E LC FF x x x x x
Nectarinia ruﬁpennis Jensen (1983) E VU FF x
Nectarinia usambarica Grote (1922)
E FF x xu x x x
Nectarinia fuelleborni Reichenow (1899)
NE FF xx
Shelley (1896) NE LC F x x x x x x x
Ploceus nicolli Sclater (1931) E EN FF x x x x
Poeoptera kenricki Shelley (1894) NE LC FF x x x x x x x x x x
Scepomycter winifredae Moreau (1938) E VU FF xx x x
Serinus whytii Shelley (1897) NE LC NF xx x
Serinus melanochrous Reichenow (1900) NE LC F xx x
Sheppardia aurantiithorax Beresford et al. (2004) EFF xx x
Grant and Mackworth-
NE VU FF xa x x
Sheppardia montana Reichenow (1907) E EN FF x
Sheppardia sharpei Shelley (1903) NE LC FF x x x x x xu x x x
Sheppardia gunningi Reichenow (1878) NE VU FF x xx
Shelley (1880) NE LC F x x x x x x x x x x
Swynnertonia swynnertoni Shelley (1906) NE VU FF x xx
Turdus helleri Mearns (1913) E CR FF x
Turdus roehli Reichenow (1905)§ E FF x x x x
Xenoperdix udzungwensis Dinesen et al. (1994) EEN FF x
Xenoperdix obscurata Fjeldsa
˚and Kiure (2003)
ˆEEN FF x
Zosterops silvanus Peter and Loveridge (1935) E EN F x
Zosterops winifredae Sclater (1935) E VU F x
Beamys hindei Thomas (1909) NE NT F x x x x x x x x x x x xu x
Cephalophus spadix True (1890) NE VU FF x x x x x
Mittermeier (1986) E FF x
Congosorex phillipsorum Stanley et al. (2005) E FF x
Crocidura desperata Hutterer et al. (1991) NE EN FF xx
Crocidura monax sensu lato Thomas (1910) NE DD FF x x x x x x x x x
Crocidura tansaniana Hutterer (1986) E VU FF x x
Crocidura telfordi Hutterer (1986) E EN FF xx
Crocidura usambarae Dippenaar (1980) E EN FF x x x
Dendrohyrax validus True (1890) NE VU FF x x x x x x x x x x x
Galagoides orinus Lawrence and Washburn (1936) E DD FF x x x x x
(continued on next page)
BIOLOGICAL CONSERVATION 134 (2007) 209–231 221
Appendix 1 – continued
Species Authority Ende-
Forest Taita North
Nguu Nguru Uluguru Ukaguru Rubeho Malundwe Mahenge Udzungwa Coastal
Galagoides zanzibaricus Matschie (1893) NE VU FF x x x x
Hylomyscus arcimontensis Carleton and Stanley (2005) NE FF x x x x x x x x x
Rungwecebus kipunji Jones et al. (2005) NE FF xx
Bergmans (1980) NE VU FF x x x x
Myosorex geata Allen and Loveridge (1927) E DD FF x
Myosorex kihaulei Stanley and Hutterer (2000) EEN FF x
Paraxerus vexillarius Kershaw (1923) NE VU FF x xu x x x x
Procolobus gordonorum Matschie (1900) E VU FF x
Rhinolophus deckenii Peters (1867) NE DD F x x x x
Rhinolophus maendeleo Kock et al. (2000) NE DD FF x x
Rhynchocyon petersi Bocage (1880) NE EN F x x x x x x x x xb
Sylvisorex howelli Jenkins (1984) E VU FF x x x x x x
Adenorhinos barbouri Loveridge (1930) NE f xx
Agama montana Barbour and Loveridge (1928) E F x x x x
Amblyodipsas teitana Broadley (1971) E F x
Aparallactus werneri Boulenger (1895) NE F x x x x xx
Atheris ceratophora Werner (1895) E F x x x
Buhoma procterae Loveridge (1922) E FF xx
Buhoma vauerocegae Tornier (1902) E FF x x
Chamaeleo deremensis Matschie (1892) E FF x x x
Chamaeleo ﬁscheri Reichenow (1887) E FF x x x x
Chamaeleo goetzei Tornier (1899) NE NF xx
Chamaeleo laterispinis Loveridge (1953) E F x
Chamaeleo oxyrhinum Klaver and Bo
¨hme (1988) E FF x x x x
Chamaeleo tempeli Tornier (1899) NE F xx
Chamaeleo tenue Matschie (1892) E F x
Chamaeleo werneri Tornier (1899) E F x x x x x
Cnemaspis barbouri Perret (1986) NE FF x x x
Cnemaspis uzungwae Perret (1986) NE FF xx
Crotaphopeltis tornieri Werner (1908) NE FF x x x x x x x
Dipsadoboa werneri Boulenger (1897) E FF x x x
Elapsoidea nigra Gu
¨nther (1888) E F x x x x x
Gastropholis prasina Werner (1904) NE F x x x
Leptosiaphos rhomboidalis Broadley (1989) E FF x
Lycophidion uzungwense Loveridge (1932) E F x
Lygodactylus conradti Matschie (1892) NE FF x x
Pasteur (1964) NE FF x x x
Lygodactylus uluguruensis Pasteur (1964) NE F x x
Lygodactylus williamsi Loveridge (1952) E FF x
Melanoseps uzungwensis Loveridge (1942) E FF x
Philothamnus macrops Boulenger (1895) NE F x x x x x x
Prosymna ornatissima Barbour and Loveridge (1928) E FF x
Prosymna semifasciata Broadley (1995) NE FF x x
Rieppeleon brevicaudatus Matschie (1892) NE F x x x x x x
Rhampholeon spinosum Matschie (1892) E FF x x
Rhampholeon moyeri Menegon et al. (2002) EFF xn x x
Rhampholeon temporalis Matschie (1892) E FF x
Tilbury and Emmrich (1996) E FF x x x x
Rhampholeon sp. nov. Matthee et al. (2004) EFF x
Rhinotyphlops nigrocandidus Broadley and Wallach (2000) EFF xx x
Rhinotyphlops gierrai Mocquard (1897) NE FF/F x xu x
Scelotes uluguruensis Barbour and Loveridge (1928) E FF x x x x
Tetradactylus udzungwensis Salvidio et al. (2004) ENF x
Thelotornis usambaricus Broadley (2001) NE F x x x x
Typhlops uluguruensis Barbour and Loveridge (1928) E F x
Typhlops usambaricus Laurent (1964) E FF x
Tornier (1900) NE FF x x x
Xyeledontophis uluguruensis Broadley and Wallach (2002) EFF x
222 BIOLOGICAL CONSERVATION 134 (2007) 209–231
Species Authority Ende-
Forest Taita North
Nguu Nguru Uluguru Ukaguru Rubeho Malundwe Mahenge Udzungwa Coastal
Afrixalus uluguruensis Barbour and Loveridge (1928) NE VU FF x x x x x x x x x
Afrixalus morerei Dubois (1986) ‘‘1985’’ E VU f x
Arthroleptides martiensseni Nieden (1910) NE EN FF x x
Arthroleptides yakusini Channing et al. (2002) EEN FF x x x x
Arthroleptis afﬁnis Ahl (1939) NE LC FF x x x x x x x
Arthroleptis reichei Nieden (1910) NE NT FF xx
Arthroleptis nikeae Poynton (2003) EEN FF x
Arthroleptis tanneri Grandison (1983) E VU FF x
Arthroleptis xenodactylus Boulenger (1909) E VU F x
Boulengerula boulengeri Tornier (1896) E LC FF x x
Boulengerula niedeni Mu
¨ller et al. (2005) EVU F x
Boulengerula taitanus Loveridge (1935) E LC F x
Boulengerula uluguruensis Barbour and Loveridge (1928) E LC F x x
Bufo brauni Nieden (1910) E EN FF x x x x x
Bufo uzunguensis Loveridge (1932) NE VU NF xx
Callulina kisiwamsitu De Sa
´et al. (2004) EFF x
Callulina kreffti Nieden (1911) ‘‘1910’’ E LC FF x x x xu x x x x x x
Churamiti maridadi Channing and Stanley (2002) ECR FF x
Hoplophryne rogersi Barbour and Loveridge (1928) E EN FF x x
Hoplophryne uluguruensis Loveridge (1925) E VU FF x x x
Hyperolius kihangensis Schiøtz and Westergaard in
EEN FF x
Hyperolius puncticulatus Pfeffer (1893) NE LC F x x x x x x x x x x x
Hyperolius minutissimus Schiøtz (1975) NE VU NF xx
Hyperolius spinigularis Stevens (1971) NE LC f x x x x x x x
Hyperolius tannerorum Schiøtz (1982) E EN FF x
Hyperolius tornieri Ahl (1931) E DD F x
Leptopelis barbouri Ahl (1929) NE VU FF x x x x x
Leptopelis uluguruensis Barbour and Loveridge (1928) E VU FF x x x x x
Leptopelis vermiculatus Boulenger (1909) NE VU FF x x x x x x x x
Leptopelis parkeri Barbour and Loveridge (1928) E VU FF x x x x x
Nectophrynoides asperginis Poynton et al. (1999) E CR NF x
Nectophrynoides cryptus Perret (1971) E EN FF x
Nectophrynoides frontierei Menegon et al. (2004) EDD FF x
Nectophrynoides laevis Menegon et al. (2004) EDD FF x
Nectophrynoides minutus Perret (1972) E EN FF xx
Nectophrynoides poyntoni Menegon et al. (2004) ECR FF x
Menegon et al. (2004) EEN FF x
Nectophrynoides tornieri Roux (1906) NE LC FF x x x x x x
Nectophrynoides vestergaardi Menegon et al. (2004) EEN FF x
Nectophrynoides viviparus Tornier (1905) NE VU F xx x x
Nectophrynoides wendyae Clarke (1988, 1989) E CR FF x
Parhoplophryne usambarica Barbour and Loveridge (1928) E CR F x
Phlyctimantis keithae Schiøtz (1974) E VU f x
Phrynobatrachus kreffti Boulenger (1909) E EN F x x x
Phrynobatrachus uzungwensis Grandison and Howell (1983) E EN FF x x x x
Probreviceps macrodactylus Nieden (1926) E VU FF x x x
Probreviceps rungwensis Loveridge (1932) NE VU FF xx
Loveridge (1925) E VU FF x
Boulenger (1883) NE LC FF x x x x x x
Barbour and Loveridge (1928) E LC FF x
Boulenger (1895) E VU FF x x x x x x x
Ahl (1924) NE LC F xxxxx
Poynton and Clarke (1999) EEN FF x
(continued on next page)
BIOLOGICAL CONSERVATION 134 (2007) 209–231 223
Appendix 1 – continued
Species Authority Ende-
Forest Taita North
Nguu Nguru Uluguru Ukaguru Rubeho Malundwe Mahenge Udzungwa Coastal
Threatened animals in the Eastern Arc that are neither endemic nor near-endemic
Sundevall (1850) No VU NF xx
Matschie (1897) No VU F x xx
Loxodonta africana Blumm enbach
No EN NF/f x x x x x X x x x
Diceros bicornis Linnaeus (1758) No CR NF/f xu xu
Panthera leo Linnaeus (1758) No VU NF/f xx
Rhynchocyon cirnei Peters (1847) No VU F/f xx x x x
Cinnyricinclus femoralis Richmond (1897) No VU F x x x
Key: Endemism – E = strictly endemi c; NE = near-endemic (also found in at least one other African ecoregion); No = not endemic. Red List – threat codes according to the IUCN red list (accessed November 2005). CR = critically endang ered, EN = endangered, VU = vulnerable,
DD = data deﬁcient, NT = near-threatened, LC = least concern, Blank = not evaluated. Forest – degree of forest dependence (FF = strictly conﬁned to forest; F = mainly forest, but also found outside; f = forest visitor; NF = non-forest species). Records – x = conﬁrmed presence;
xa = record from the Andersen collection in Copenhagen; xb = recent camera trap photos cast doubt on taxonomy of these records; xn = new species in the pro cess of being described, but retained within the original taxon for the purposes of this paper; xu = record uncertain
(and not used for analysis); ** = species not formally described; *** = updated taxonomy in Roy et al. (1997) is not followed here; **** = taxonomy updated by Bowie et al. (2004); § = taxonomy updated by Bowie et al. (2004);ˆ= taxonomy updated by Bowie and Fjeldsa
Distributional notes on some Eastern Arc near-endemic species that are found elsewhere in Africa.. Oriolus chlorocephalus = Southern Malawi into Mozamb ique, Sheppardia lowei, xa = Uluguru record from Andersen collection, Stactolaema olivacea = South Africa, Hirundo
atrocaerulea = Udzungwa, Southern Rift, Uganda, South Africa, Malawi, Myonycteris relicta = also Haroni-Rusitu in Zimbabwe, Lygodactylus gravis = Maji Kununua (Mkomazi), Urocotyledon wolterstorfﬁ = possible record from Arusha.
224 BIOLOGICAL CONSERVATION 134 (2007) 209–231
Appendix 2. Distribution of endemic and near endemic trees across the forests of the Eastern Arc Mountains
Uluguru Ukaguru Rubeho Malundwe Udzungwa Maheng e Near endemic
Anisophyllea obtusifolia Engl. and Brehm.
Annonaceae gen. nov. = Ede 65
Anonidium usambarense R.E. Fries
Enantia kummeriae Engl. and Diels
Greenwaydendron suaveolens (Engl. and Diels) Verdc. subsp. usambaricum Verdc.
Isolona heinsenii Engl. and Diels
1 1 Also coastal
Lettowianthus stellatus Diels U
1 1 1 1 1 Also coastal
Polyceratocarpus schefﬂeri Engl. and Diels
Uvariodendron gorgonis Verdc.
1 1 1 1 Also coastal
Uvariodendron oligocarpum Verdc.
Uvariodendron pycnophyllum (Diels) R.E. Fries
Uvariodendron usambarense R.E. Fries
Polyscias stuhlmannii Harms
Schefﬂera lukwangulensis (Tennant) Bernardi
Platypterocarpus tanganyikensis Dunkley and Brenan
Hirtella megacarpa R. Grah.
Allanblackia stuhlmannii (Engl.) Engl.
111111 1 1
Allanblackia ulugurensis Engl.
Garcinia semseii Verdc.
11 1 1
Mammea usambarensis Verdc.
Diospyros kabuyeana F. White U1 1 1 1 Also coastal
Diospyros occulta F. White
11 1 Also coastal
Diospyros sp. aff. amaniensis Guerke
Croton dictyophlebodes A.R.-Sm.
Drypetes usambarica (Pax) Hutch.
1 1 1 1 1 1 Also coastal
Macaranga conglomerata Brenan
Sibangea pleioneura A.R.-Sm.
Angylocalyx braunii Harms U
11 1 Also coastal
Cynometra brachyrrachis Harms
1 Also coastal
Cynometra engleri Harms
Cynometra longipedicellata Harms
Cynometra sp. A
Cynometra sp. B
Cynometra ulugurensis Harms
Englerodendron usambarense Harms
Isoberlinia schefﬂeri (Harms) Greenway
1 1 1 1 1 1 1 Also coastal
Millettia elongatistyla Gillett
Newtonia paucijuga (Harms) Brenan
1 1 1 1 Also coastal
Pterocarpus mildbraedii Harms subsp. usambarensis (Verdc.) Polhill
Scorodophloeos ﬁscheri (Taub.) J. Le
1 1 1 1 1 1 Also coastal
Zenkerella capparidacea (Taub.) J. Le
Zenkerella egregia J. Le
111 Also coastal
Zenkerella perplexa Temu
(continued on next page)
BIOLOGICAL CONSERVATION 134 (2007) 209–231 225
Appendix 2 – continued
Uluguru Ukaguru Rubeho Malundwe Udzungwa Mahenge Near endemic
Dasylepis integra Warb.
111 Also Mbulu
Casearia engleri Gilg
Beilschmidia kweo (Mildbr.) Robyns and Wilczek
11 1 1
Lijndenia brenanii (A. and R. Fernandes) Jacq. Fe
Lijndenia greenwayii (Brenan) Borhidi
Memecylon sp. A
Memecylon teitense Wickens
Cephalosphaera usambarensis (Warb.) Warb.
1 1 1 1 1 Also Shimba Hills
Syzygium micklethwaitii Verdc.
Ouratea schefﬂeri Engl. and Gilg
Ouratea schusteri Gilg ex Engl.
11 1 1
Octoknema orientalis Mildbr.
Pittosporum goetzei Engl.
Calodendrum eickii Engl.
Allophylus melliodorus Radlk.
Pancovia sp. B 1 Also North Malawi
Placodiscus amaniensis Radlk.
Placodiscus pedicellatus F.G. Davies
Neohemsleya usambarensis Pennington
Omphalocarpum strombocarpum Y. B. H a r v. a n d J. C . L ov e tt
Pouteria pseudoracemosa (J.H. Hemsl.) L. Gautier
Cola schefﬂeri K. Schum.
Leptonychia usambarensis K. Schum.
1 111111 1
Grewia goetzeana K. Schum.
11 Also coastal
Vitex amaniensis Pieper
Totals 8 0 1 27 40 6 25 26 4 0 4 36 5
Notes on habitat associations: § = submontane forest; U= dry lowland forest; = lowland forest; *= montane forest; k= dry montane forest; à= upper montane forest.
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