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Priority Primate Areas in Tanzania
T IM R.B. DAVENPORT,KATARZYNA N OWAK and A NDREW P ERKIN
Abstract Priority Primate Areas are identified in Tanzania,
mainland Africa’s most important country for conservation
of primates, on the basis of occupancy by globally rare, Red-
Listed and range-restricted primate species and subspecies.
We provide a comprehensive list and regional assessment of
Tanzania’s primate taxa, using IUCN Red List criteria, as
well as the first national inventory of primates for 62 sites.
The Priority Primate Areas, encompassing 102,513 km
2
,
include nine national parks, one conservation area, seven
game reserves, six nature reserves, 34 forest reserves and five
areas with no official protection status. Primate species were
evaluated and ranked on the basis of irreplaceability and
vulnerability, using a combination of established and ori-
ginal criteria, resulting in a primate Taxon Conservation
Score. Sites were ranked on the basis of summed primate
scores. The majority (71%) of Priority Primate Areas are also
Important Bird Areas (IBAs), or part of an IBA. Critical
subsets of sites were derived through complementarity
analyses. Adequate protection of just nine sites, including
six national parks (Kilimanjaro, Kitulo, Mahale, Saadani,
Udzungwa and Jozani-Chwaka Bay), one nature reserve
(Kilombero) and two forest reserves (Minziro and
Mgambo), totalling 8,679 km
2
, would protect all 27 of
Tanzania’s primate species. The addition of three forest
reserves (Rondo, Kilulu Hill and Ngezi) and two game
reserves (Grumeti and Biharamulo), results in a list of 14
Priority Primate Areas covering 10,561 km
2
(1.1%of
Tanzania’s total land area), whose conservation would
ensure the protection of all 43 of Tanzania’s species and
subspecies of primates.
Keywords Conservation status, primates, priority sites,
protected areas, Tanzania
This paper contains supp lementary material that can be
found online at http://journals.cambridge.org
Introduction
S
ince the mid 1990s a variety of priority setting concepts
have been designed to guide policy and help reap
maximum benefit in a world of limited conservation
resources (Balmford, 2002; Caro, 2010; Gauthier et al.,
2010). Often employing different taxa and/or criteria, these
have been either coarse-scale and global in reach (e.g. Global
Biodiversity Hotspo ts, Myers et al., 2000; Centres of Plant
Diversity, WWF & IUCN 199 4–1997; Endemic Bird Areas,
Stattersfield et al., 1998 ), or more fine-scale and regional,
(e.g. Important Bird Areas, Fishpool & Evans, 2001; Impor-
tant Plant Areas, Anderson, 2002 , Plantlife International,
2004; Important Mammal Areas, Linzey, 2002; Prime
Butterfly Areas, van Swaay & Warren, 2003). Others (e.g.
Key Biod iversity Areas) are sites of global importance for
conservation of biodiversity and yet aimed at the site-scale
(Langhammer et al., 2007), with the irreplaceability and
vulnerability of species the main criteria.
Tanzania is widely regarded as being the most important
country in mainland Africa for biodiversity and biological
endemism, with the continent’s highest mountain, deepest
lakes and two globally significant biodiversity hotspots, the
Eastern Arc Mountains and the Albertine Rift (Burgess
et al., 2004a, 2004b). However, Tanzania has the second
highest rate of forest loss in sub-Saharan Africa (FAO, 2005)
and, despite considerable conservation investment and a
large amount of land nominally under protection, natural
habitats and biodiversity continue to be lost at a significant
rate (e.g. Rovero et al., 2012). For a developing nation of
such global natural resource importance, priority setting is
an essential conservation tool.
A range of general national-level priority setting analyses
has been carried out in Tanzania. The African Mammal
Databank undertook an environmental suitability and
species occurrence analysis (Boitani et al., 1999), although
extrapolations were based on species habitat suitability
rather than species occupancy. No distinction was made
at the subspecies level, and the extent and number of
occupied protected areas was overestimated. The European
Commission subsequently used these data to assess 31 pro-
tected areas in Tanzania for mammal, bird, amphibian and
habitat irreplaceability (Hartley et al., 2007), and IUCN’s
Gap Analysis evaluated whether protected areas safeguard
biodiversity and how much biodiversity falls outside them
(Langhammer et al., 2007). More specifically, the Alliance
for Zero Extinction (Ricketts et al., 2005) aimed to identify
sites that contain at least 95% of a known population of one
or more Critically Endangered or Endangered species. In
Africa, 76 sites and 122 species were identified and Tanzania
had the most (eight) Alliance for Zero Extinction sites,
seven of which are based on amphibians. However, because
of the nature of the Alliance for Zero Extinction process
there is a bias towards sites that have been better
investigated, and thus the list for Tanzania is neither topical
TIM DAVENPORT (Corresponding author) Wildlife Conservation Society, P.O.
Box 922, Zanzibar, Tanzania. E-mail: tdavenport@wcs.org
K
ATARZYNA NOWAK Udzungwa Elephant Project, Iringa, Tanzania
A
NDREW PERKIN c/o Tanzania Forest Conservation Group, Dar es Salaam,
Tanzania
Received 14 July 2012. Revision requested 26 October 2012.
Accepted 16 November 2012.
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nor complete. Currently, the Critical Ecosystem Partnership
Fund is identifying Key Biodiversity Areas across the
Eastern Afromontane Biodiversity Hotspot, including in
Tanzania, but this work is not yet complete (BirdLife, 2012).
At the species level there have been only two national
analyses, the 77 Important Bird Areas of Tanzania (Baker &
Baker, 2002) and a partial identification of the nation’s
Important Plant Areas (McClean et al., 2006).
Primates are one of the most threatened and charismatic
groups. The taxon is largely forest-dependent and Tanzania
hosts eight endemic species. This represents 29% of its total
primate species, a figure second only to Madagascar.
Usually, it is the rare primate species that are less well
studied, and this rarity is attributed to ecological special-
ization (Doherty & Harcourt, 2004). In Tanzania, however,
the two rarest species are not obligate specialists (Davenport
et al., 2010; Nowak & Lee, 2011) and as little is known about
their distribution and conservation status as about that of
the more common species. With the exception of a species
list recently constructed as part of a photographic atlas of
East African primates (De Jong & Butynski, 2012) and a few
site-specific primate lists (e.g. Rovero et al., 2009), Tanzania
lacks a complete annotated list of its species (and
subspecies/races) of diurnal and nocturnal primates, or a
nationwide dataset of primate distribution and status. We
provide here the fi rst complete annotated list of Tanzanian
primates, including diurnal and nocturnal species and
subspecies. We also present a novel concept synonymous
with Important Bird Areas, which we have called Priority
Primate Areas, based on our compilation and comprehen-
sive nationwide analysis of distributions and status. We
identify these Priority Primate Areas to facilitate a focus on
the range-restricted and threatened primate species and
subspecies that will most benefit from conservation at the
site-level. The intention is to quantify at a national scale the
immediacy and precedence of threats to a group that has
both considerable conservation relevance and charismatic
appeal, thus identifying conservation gaps, providing ‘watch
lists’ of priority sites and taxa using globally consistent
criteria (Langhammer et al., 2007), and ensuring there is an
objective, empirical and ranked plan to protect all of
Tanzania’s primates.
Specifically, we aim to (1) identify major populations and
subpopulations of all primate species and subspecies in
Tanzania, (2) rank species and subspecies according to their
irreplaceability, vulnerability and conservation status (and
anticipating future taxonomic change), (3) identify priority
sites for conservation of primates on the basis of number
of species present and rank, (4) provide a minimum critical
list of sites whose protection would ensure the conservation
of all of Tanzania’s primates, (5) complement other taxa-
specific analyses such as Important Bird Areas, (6) identify
gaps in knowledge and conservation at a national scale, thus
limiting duplication of conservation effort, and (7) provide
a concept (Priority Primate Areas) that could be adopted
internationally to prioritize implementation of the con-
servation of primate taxa. Primates are often flagship species
(Caro, 2010); by identifying and quantifying priority sites for
primates in Tanzania we hope to ameliorate resource al-
location problems facing government and conservation
organizations (Rondinini et al., 2006; Wilson et al., 2006)
and focus conservation attention on Tanzania’s unique
primate fauna and the habitats in which they dwell.
Methods
Species list
A full species list of the diurnal and nocturnal primates of
Tanzania was drawn up (Table 1). We followed the taxo-
nomy of Grubb et al. (2003), and referred to Groves (2007)
for Lophocebus johnstoni, Kingdon (1997) and Groves
(2005) for Cercocebus sanjei, Davenport et al. (2006) for
Rungwecebus kipunji,Groves(2001, 2005) for Chlorocebus
pygerythrus, and Rahm (1970), Napier (1985) and Kingdon
(1997) for Colobus angolensis sharpei. Primate taxonomy is
developing, occasionally contentious, and compounded by
concepts such as superspecies and clines, and by personal
interpretation. It is not our intention here to enter into
taxonomic debate. Tanzanian primates also experience high
levels of hybridization, such as between the baboons Papio
anubis and Papio cynocephalus on the Tanzania–Kenya
border (Alberts & Altmann, 2001) and the several inter- and
intraspecific Cercopithecine hybrid zones (Detwiler et al.,
2005), including Cercopithecus mitis albogularis and
Cercopithecus mitis stuhlmanni in Ngorongoro and Lake
Manyara, and Cercopithecus ascanius schmidti and
Cercopithecus mitis doggetti in Gombe National Park
(Detwiler, 2002). Although we recognize the presence of
hybrids we did not include them in the analysis. We
recognize two forms of Galagoides rondoensis, based on
recent data (Perkin et al., unpubl. data).
Species scores
For each species and subspecies IUCN (2012b) proscriptions
were applied. Following IUCN (2003) we also made
assessments of the extinction risk of Tanzanian national
populations of species and subspecies, as such evaluations
were previously lacking. We thus assigned regional Red List
categories to all Tanzanian taxa if the national status mer-
ited a classification higher than the global status (Table 1).
Nationwide primate distributions were comp iled from the
literature, online databases, local expert knowledge and
personal observations from our combined experience in
Tanzania. Databanks included the IUCN Red List (IUCN,
2012b),thePrimateInformationNetworkfactsheets(Primate
Info Net, 2011), the African Mammals Databank (1999),
2 T. R. B. Davenport et al.
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TABLE 1 List of primate species and subspecies in Tanzania, with their Red List status at the global (IUCN, 2012b) and national levels (see text for sources). Endemic taxa are in bold (note that
national and global assessments are the same when a taxon is endemic).
Genus Species Subspecies Common name
Red List status
1
(criteria
2
)
Global sp. Global subsp. National sp. National subsp.
Hominidae
Pan troglodytes schweinfurthii Eastern robust chimpanzee EN EN EN
Cercopithecidae
Cercocebus sanjei Sanje mangabey EN EN
Cercopithecus ascanius schmidti Schmidt’s red-tailed monkey LC LC NT*
mitis monoides Tanzania Sykes’ monkey LC LC LC LC
moloneyi Moloney’s white-collared monkey LC LC
doggetti Doggett’s blue monkey LC LC
albogularis Zanzibar Sykes’ monkey LC LC
Chlorocebus pygerythrus hilgerti Hilgert’s vervet LC LC LC
nesiotes Pemba vervet VU(B1a,bi,iii)*
rufoviridis Reddish-green vervet LC
tantalus Tantalus monkey LC
Erythrocebus patas baumstarki Serengeti patas monkey LC VU(B1a,b,C2a,D1)* VU(B1a,b,C2a,D1)*
Lophocebus ugandae Uganda grey-cheeked mangabey LC E(B1a,biii,B2a,b)*
Papio anubis Olive baboon LC LC
cynocephalus cynocephalus Yellow baboon LC LC LC LC
kindae Kinda yellow baboon LC LC
Rungwecebus kipunji Kipunji CR CR
Colobidae
Colobus angolensis palliatus Peters’ Angola colobus LC LC LC LC
sharpei Sharpe’s Angola colobus VU(B1a,b,c)* VU(B1a,b,c)*
ruwenzorii Adolf Friedrichs’s Angola colobus VU NE
subsp. nov. Nkungwe Angola colobus DD NE
guereza caudatus Mt Kilimanjaro guereza LC LC LC VU(B1a,b)*
matschei Mau Forest guereza LC NT*
Procolobus rufomitratus tephrosceles Eastern red colobus LC EN EN
gordonorum Udzungwa red colobus EN EN
kirkii Zanzibar red colobus EN EN
Galagidae
Galago senegalensis braccatus Kenya lesser galago LC LC LC LC
sotikae Uganda lesser galago LC LC
moholi Southern lesser galago LC LC
Priority Primate Areas in Tanzania 3
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the Tanzania Mammal Atlas Project (2013), the Pictorial
Guide to Living Primates (2013), Tanzania National
Parks websites and grey literature, although these mainly
provided extent of occurrence estimates rather than occu-
pancy.
Site list
A list of discrete sites in which species occur was compiled
(Fig. 1). This followed the Alliance for Zero Extinction
definition (Ricketts et al., 2005): ‘an area with a definable
boundary within which the character of habitats, biological
communities, and/or management issues have more in
common with each other than they do with those in ad-
jacent areas.’ Thus, sites such as Tongwe East (Forest
Reserve) and Ugalla River (Game Reserve), and Kitulo
(National Park) and Mt Rungwe (Nature Reserve) were
treated separately, although contiguous, because of their
different protected area status and hence management.
Protected area data came from Protected Planet (2013), the
Tanzania Ministry of Natural Resources and Tourism,
and Tanzania National Parks. Areas to which species were
introduced (Rubondo Island for Pan troglodytes and
Colobus guereza, and Ngezi Forest, Pemba, for Procolobus
kirkii), were not included although we recognize that these
sites may contribute to species’ conservation and genetic
and behavioural diversity.
Taxon conservation score
To rank sites it was first necessary to assign conservation
scores to each taxon so that a summed score based on taxon
presence could be applied. The taxon conservation score
is a sum of a taxon’s irreplaceability and vulnerability.
For irreplaceability we modified the index used by the
European Community (Hartley et al., 2007), calculating 1/n
(maximum 5 1), where n 5 the number of sites in which a
taxon occurs. To this site occupancy score we added a value
reflecting the percentage of a taxon’s total range that falls
within Tanzania (maximu m 5 2.5; Table 2); this gives a
proxy for percentage of the total population within
Tanzania, data that are unavailable for most taxa. Thus
taxon irreplaceability score 5 (1/n) + (% range score). We
calculated a taxon vulnerability score using global Red List
data (IUCN, 2012b) and our recommended Tanzanian Red
List status (maximum 5 4; Table 3), and the current popu-
lation trend for each taxon. The latter was quantified
according to whether the Tanzanian population is decreas-
ing (0.5), stable or assumed stable (0), increasing (−0.5)or
unknown (−0.25). Thus vulnerability score 5 Red List
score + population trend score. Summing the scores of
taxon irreplaceability and vulnerability produces an overall
taxon conservation sco re (maximum total 5 8).
Table 1 (Cont.)
Genus Species Subspecies Common name
Red List status
1
(criteria
2
)
Global sp. Global subsp. National sp. National subsp.
Galagoides cocos Kenya coast galago NT EN
demidoff anomurus Demidoff’s galago LC VU VU
granti Mozambique galago LC LC
orinus Mountain galago NT NT*
rondoensis CR CR
subsp. A Rondo galago CR CR
subsp. B Rondo galago CR CR
sp. nov. Rungwe galago EN(B1abi,ii)* EN(B1abi,ii)*
thomasi Thomas’ galago LC VU
zanzibaricus udzungwensis Matundu galago LC LC LC LC
zanzibaricus Zanzibar galago EN EN
Otolemur crassicaudatus montieri Large-eared greater galago LC LC
garnettii garnettii Zanzibar small-eared galago LC LC LC LC
lasiotis White-tailed small-eared galago LC VU
panganiensis Pangani small-eared galago LC LC
1
CR, Critically Endangered; EN, Endangered; VU, Vulnerable; NT, Near Threatened; LC, Least Concern; NE, Not Evaluated
2
IUCN (2003, 2012a)
*Recommended change, or assessed in this study
4 T. R. B. Davenport et al.
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Site scores
A matrix was compiled of the 62 sites identified (Fig. 1) and
occupancy by primates (Supplementary Table S1). The
taxon conservation scores for primates at each site were
summed, producing total site scores for nocturnal, diurnal
and all (combined nocturnal and diurnal) species and all
taxa (Supplementary Tables S2 &S3, respectively). Sites
were then ranked according to their taxon scores: all species,
nocturnal species, diurnal species, all taxa, nocturnal taxa
and diurnal taxa. To derive a ranked list of Priority Primate
Areas two simple complementarity analyses were per-
formed, one by taxa frequency and the other by site rank.
For the former we began with the site in each category that
had the highest number of taxa and then added the next site
that would add more taxa. This was continued until a list of
Priority Primate Areas for species and taxa had been
produced. For the latter we began with the site that ranked
highest and subsequent complementary sites were added
consecutively from the site rank list that added more species
(Howard et al., 2000; Brugiere, 2012).
Results
We recognize 27 species of primates in Tanzania (15 diurnal
and 12 nocturnal species) and 43 taxa (26 diurnal and 17
nocturnal) including subspecies (Table 1). Of these, four are
endemic diurnal primate species (C. sanjei, R. kipunji,
Procolobus gordonorum and P. kirkii), four are endemic
nocturnal species (Galagoides orinus, G. rondoensis,
FIG. 1 Location of the 62 sites considered in the ranking analysis of Priority Primate Areas.
TABLE 2 Scores assigned to the percentage of the total range of a
primate species or subspecies that falls within Tanzania, for the
calculation of the taxon conservation score (see text for details).
% of range Species Subspecies
100 2.50 1.25
.50 2.00 1.00
20–50 1.50 0.75
5–20 1.00 0.50
,5 0.50 0.25
TABLE 3 Scores assigned to the global Red List status (IUCN, 2012b)
or our recommended Tanzanian Red List status of primate species
or subspecies (Table 1), for the calculation of the taxon
conservation score (see text for details). The highest possible
score for each taxon was applied.
Red List status
Global Tanzania
Species Subspecies Species Subspecies
Critically
Endangered
42 10.5
Endangered 3 1.5 0.75 0.375
Vulnerable 2 1 0.5 0.25
Near Threatened 1 0.5 0.25 0.125
Least Concern 0 0 0 0
Priority Primate Areas in Tanzania 5
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Galagoides sp. nov. and Galagoides zanzibaricus), and four
are endemic diurnal subspecies (Chlorocebus pygerythrus
nesiotes, Erythrocebus patas baumstarki, Colobus angolensis
subsp. nov. and Colobus guereza caudatus). The list includes
two species that are categorized on the IUCN Red List
as Critically Endangered, five as Endangered, two as Near
Threatened and 18 as Least Concern (IUCN, 2012b). At
the subspecies level two are categorized as Critically
Endangered, three as Endangered, four as Vulnerable, one
as Data Deficient and 19 as Least Concern (IUCN, 2012b). At
the national level two species are categorized as Critically
Endangered, eigh t as Endangered, three as Vulnerable,
two as Near Threatened and 11 as Least Concern, and at
national subspecies level, two as Critically Endangered,
one as Endangered, four as Vulnerable, one as Near
Threatened, two as Not Evaluated and 13 as Least Concern
(IUCN, 2012b). We assigned putative Red List categories
(IUCN, 2003) for 11 taxa; the relevant justifications are given
in Table 1.
The comparative rankings of each taxon are illustrated in
Tables 4–5. The taxon conservation scores produced the
same top six primates irrespective of whether the analysis
was at the species or taxon level. These were R. kipunji,
G. rondoensis, C. sanjei, P. gordonorum, P. kirkii and
Galagoides sp. nov. Of the 62 sites identified as being of
importance for conservation of primates 57 are officially or
nominally protected (nine national parks; six nature
reserves; 34 forest reserves, including four proposed nature
reserves; seven game reserves; and one conservation area).
The other five sites are either privately managed or have no
protection status.
On application of the taxon conservation scores to the 62
sites, a matrix was drawn up illustrating the Priority Primate
Areas across Tanzania according to all diurnal and
TABLE 4 The comparative ranking of each primate species based on
the taxon conservation score (maximum 5 8; see text for details).
Rank Species Score
1 R. kipunji 7.33
2 G. rondoensis 7.13
3 C. sanjei 6.50
4 P. kirkii 6.33
5 P. gordonorum 6.24
6 G. sp. nov. 6.20
7 P. troglodytes 4.17
8 G. orinus 4.07
9 G. cocos 3.50
10 L. ugandae 3.25
11 G. zanzibaricus 2.54
12 G. thomasi 2.50
13 P. rufomitratus 2.45
145 E. patas 2.00
145 G. demidoff 2.00
16 G. moholi 1.75
17 G. granti 1.60
185 C. angolensis 1.53
185 O. garnettii 1.53
20 C. mitis 1.52
21 C. guereza 1.25
22 G. senegalensis 1.11
23 C. ascanius 1.08
245 P. cynocephalus 1.02
245 C. pygerythrus 1.02
26 O. crassicaudatus 0.53
27 P. anubis 0.11
TABLE 5 The comparative ranking of each primate taxon (i.e.
species or subspecies) based on the taxon conservation score
(maximum 5 8 ; see text for details).
Rank Taxon Score
1 R. kipunji 7.33
2 C. sanjei 6.50
35 P. kirkii 6.33
35 P. gordonorum 6.25
5 G. rungwe sp. nov. 6.20
6 G. zanzibaricus zanzibaricus 4.50
7 P. troglodytes schweinfurthi 4.42
8 G. orinus 4.07
9 G. rondoensis subsp. nov. B 3.83
10 G. rondoensis subsp. nov. A 3.70
115 C. pygerythrus nesiotes 3.50
115 G. cocos 3.50
13 L. ugandae 3.25
14 C. angolensis subsp. nov. 2.75
15 P. rufomitratus tephrosceles 2.70
16 G. zanzibaricus udzungwensis 2.55
175 C. angolensis ruwenzorii 2.50
175 G. thomasi 2.50
195 E. patas baumstarcki 2.25
195 G. demidoff anomurus 2.25
21 O. garnettii lasiotis 2.00
225 C. angolensis sharpei 1.75
225 C. guereza caudatus 1.75
225 G. senegalensis sotikae 1.75
225 G. moholi 1.75
26 C. guereza matschiei 1.63
27 G. granti 1.60
28 O. garnettii panganiensis 1.28
29 P. cynocephalus kindae 1.25
30 C. ascanius schmidti 1.08
31 P. cynocephalus cynocephalus 1.03
32 C. mitis doggetti 1.00
33 O. garnettii garnettii 0.95
34 C. pygerythrus hilgerti 0.89
355 C. mitis monoides 0.88
355 G. senegalensis braccatus 0.88
37 C. mitis moloneyi 0.84
38 C. pygerythrus rufoviridis 0.83
39 C. angolensis palliatus 0.79
40 C. mitis albogularis 0.78
41 C. pygerythrus tantalus 0.54
42 O. crassicaudatus montieri 0.53
43 P. anubis 0.11
6 T. R. B. Davenport et al.
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nocturnal species and all diurnal and nocturnal taxa
(Supplementary Tables S2–S3). When all primates are con-
sidered, either at the species level or species and subspecies
level, the sites that emerged to be most important are
Kilombero Nature Reserve, Udzungwa Mountains National
Park and Uzungwa Scarp Forest Reserve, with Kitulo
National Park and Mt Rungwe Nature Reserve in joint
fourth place (Fig. 2). If diurnal and nocturnal primates are
treated separately, Mahale Mountains National Park is
placed fourth for diurnal taxa (Fig. 3). The nocturnal
primates produced a different assessment. At both the
species and all taxa levels, Saadani National Park, Pande
Game Reserve, and Pugu/Kazimzumbwe, Chitoa, Litipo,
Rondo, Ruawa and Ziwani Forest Reserves all illustrated the
1–10
11–20
21–30
31–62
1–10
11–20
21–30
31–62
(a)
(b)
FIG. 2 Top-ranking sites for (a) all
primate species and (b) all taxa
(i.e. species and subspecies). All
species: 1, Kilombero Nature
Reserve*†; 2, Udzungwa Mts
National Park†; 3, Uzungwa Scarp
Forest Reserve†; 45, Mt Rungwe
Nature Reserve*†; 45, Kitulo
National Park*†; 6, Saadani
National Park; 7, Mahale National
Park; 85, Pande GR*†; 85, Pugu/
Kazimzumbwe Forest Reserve*†;
10, Magombera†. All taxa: 1–5,as
above; 6, Mahale National Park; 7,
Magombera; 8, Jozani-Chwaka
Bay National Park†; 9, Minziro
Forest Reserve; 105, Kiwengwa-
Pongwe Forest Reserve†; 105, Uzi
and Vundwe Islands†.
*Critically Endangered species;
†endemic species present.
Priority Primate Areas in Tanzania 7
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importance of Tanzania’s coastal forests for galago taxa
(Fig. 4).
To derive an applied and ranked list of priority areas for
conservation of primates, two sets of Priority Primate Areas
were determined based on the two complementarity
analyses. For both the taxa frequency analysis and the site
rank analysis, a list of nine sites (with possible alternatives)
was identified for the 27 primate species, and 14 sites (with
alternatives) for the 43 primate taxa (Tables 6–7). Seven
Priority Primate Areas were identified separately for the 15
diurnal species, eight for the 12 nocturnal species, 11 for the
26 diurnal taxa, and 10 for the 17 nocturnal taxa.
1–10
11–20
21–30
31–62
1–10
11–20
21–30
31–62
(a)
(b)
FIG. 3 Top-ranking sites for
(a) diurnal primate species and
(b) taxa (i.e. species and
subspecies). Diurnal species: 1,
Kilombero Nature Reserve*†; 25,
Udzungwa Mts National Park†;
25, Uzungwa Scarp Forest
Reserve†; 4 , Mahale National
Park; 55, Mt Rungwe Nature
Reserve*†; 5 5, Kitulo National
Park*†; 7, Magombera†; 8, Gombe
National Park; 9, Jozani-Chwaka
Bay National Park†; 105,
Kiwengwa-Pongwe Forest
Reserve†; 105 , Uzi/Vundwe
Islands†. Diurnal taxa: 1 –55 ,as
above; 7, Gombe National Park;
8, Magombera; 9, Jozani-Chwaka
Bay National Park; 10, Minziro
Forest Reserve.
*Critically Endangered species;
†endemic species present.
8 T. R. B. Davenport et al.
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Discussion
Given that endemism of primates in Tanzania is 29.6%
at species level and 28.6% at species and subspecies level
combined, with the highest number of threatened primate
taxa in mainland Africa, the country is of global con-
servation significance. However, Tanzania still faces
substantial challenges, with a growing human population,
ever pressing needs for development and growth, and
an increasingly fragmented and human-dominated land-
scape. Prioritization of conservation interventions is thus
of particular importance for a nation with such a
responsibility. Because of their diversity, broad appeal and
taxonomic tractability, birds have often been used as
1–10
11–20
21–30
31–62
1–10
11–20
21–30
31–62
(a)
(b)
FIG. 4 Top-ranking sites for
(a) nocturnal primate species and
(b) taxa (i.e. species and
subspecies). Nocturnal species: 1,
Saadani National Park; 25, Pande
GR*†; 25 Pugu/Kazimzumbwe
Forest Reserve*†; 45, Ruawa
Forest Reserve; 45, Chitoa Forest
Reserve; 45, Litipo Forest
Reserve*†; 45, Rondo Forest
Reserve*†; 45, Ziwani Forest
Reserve*†; 9, Udzungwa Mts
National Park†; 10, Kilombero
Nature Reserve†. Nocturnal taxa:
1, Udzungwa Mts National Park; 2,
Saadani National Park; 3,
Kilombero Nature Reserve; 45,
Amani Forest Reserve; 45,
Mkungwe Forest Reserve; 45,
Nilo Nature Reserve; 7, Selous GR;
85, Pande GR; 85, Pugu/
Kazimzumbwe Forest Reserve;
105, Mt Rungwe Nature Reserve†
105, Kitulo National Park†.
*Critically Endangered species;
†endemic species present.
Priority Primate Areas in Tanzania 9
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effective indicators of biodiversity and habitat health
(Howard et al., 1998), although cross-taxon congruence
has been less reliable, in part because of birds’ greater
mobility (Tsushabe et al., 2006). Despite their charismatic
qualities and evolutionary proximity to humans, primates
have been less of a focus for priority setting and less often
used as surrogate taxa for broader conservation needs. This
priority setting for primates is the first such national
analysis, and offers an objective, quantifiable protocol for
the identification of Priority Primate Areas. We have
developed this method both to provide the means to
replicate the process of prioritizing sites important for
primates in other countries and to help focus national
conservation priorities in Tanzania.
There were a number of challenges, and compromises
made, in design ing this system, the foremost being how best
TABLE 6 Minimum critical set of Priority Primate Areas, based on
complementarity analyses (see text for further details), for all,
diurnal and nocturnal species. The sites are those that would at a
minimum need to be protected to conserve at least one population
of each of Tanzania’s primate species.
Rank Area*
No. of
species
All species
1 Kilombero NR 10
2 Udzungwa Mts NP 2
3 Kitulo NP or Mt Rungwe NR 1
4 Saadani NP 1
5 Mahale Mountains NP 5
6 Jozani-Chwaka Bay NP 1
7 Minziro FR 4
8 Mgambo FR 1
9 Kilimanjaro NP 2
Total 27
Diurnal species
1 Kilombero NR 6
2 Udzungwa Mts NP or
Uzungwa Scarp FR
1
3 Mahale Mts NP 3
4 Gombe Stream NP 1
5 Jozani-Chwaka Bay NP 1
6 Minziro FR 1
7 Arusha NP or Kilimanjaro NP 2
Total 15
Nocturnal species
1 Sadaani NP 4
2 Chitoa FR or Litipo FR or Rondo
FR or Ruawa FR or Ziwani FR
1
3 Udzungwa Mts NP 1
4 Kilombero NR 1
5 Selous 1
6 Kitulo NP or Mt Rungwe NR 1
7 Kilulu Hill FR or Mgambo FR 1
8 Minziro FR 2
Total 12
*NR, Nature Reserve; NP, National Park; FR, Forest Reserve
TABLE 7 Minimum critical set of Priority Primate Areas, based on
complementarity analyses (see text for further details), for all,
diurnal and nocturnal taxa (i.e. species and subspecies). The sites
are those that would at a minimum need to be protected to
conserve at least one population of each of Tanzania’s primate taxa.
Rank Area*
No. of
taxa
All taxa
1 Kilombero NR 10
2 Udzungwa Mts NP 3
3 Kitulo NP or
Mt Rungwe NR
2
4 Mahale Mountains NP 8
5 Jozani-Chwaka Bay NP 5
6 Minziro FR 5
7 Saadani NP 1
8 Chitoa FR or Litipo FR or
Rondo FR or Ruawa FR
or Ziwani FR
1
9 Kilimanjaro NP 3
10 Mgambo FR 1
11 Kilulu Hill FR 1
12 Grumeti GR 1
13 Biharamulo GR 1
14 Ngezi FR 1
Total 43
Diurnal taxa
1 Kilombero NR 6
2 Udzungwa Mts NP or
Uzungwa Scarp FR
2
3 Mahale Mts NP 6
4 Kitulo NP or Mt Rungwe NR 1
5 Gombe Stream 1
6 Jozani-Chwaka Bay NP 3
7 Minziro FR 2
8 Tongwe East GR 1
9 Arusha NP or
Kilimanjaro NP
2
10 Grumeti GR 1
11 Ngezi FR 1
Total 26
Nocturnal taxa
1 Udzungwa Mts NP 4
2 Sadaani NP 1
3 Kilombero NR 1
4 Selous GR 2
5 Kitulo NP or Mt Rungwe NR 1
6 Chitoa FR or Litipo FR or
Rondo FR or Ruawa FR or
Ziwani FR
1
7 Kilulu Hill FR 2
8 Jozani NP or Masingini FR
or Uzi-Vundwe
or Kiwengwa-Pongwe
2
9 Minziro FR 2
10 Biharamulo GR 1
Total 17
*NR, Nature Reserve; NP, National Park; FR, Forest Reserve; GR, Game
Reserve
10 T. R. B. Davenport et al.
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to define a site. We concluded that an area must have a
definable boundary within which the character of habitats,
biological communities, and/or management issues have
more in common with each other than they do with those in
adjacent areas. Conservation must be pragmatic. However,
it is clear that prioritization of forest sites can change as
lesser known areas are investigated (Burgess et al., 2007;
Davenport et al., 2007). That notwithstanding, the corre-
lation between site rankings and degree of survey effort
(Doggart et al., 2006) is less of an issue with smaller
taxonomic groups such as primates.
There are a number of sites in Tanzania where primate
species have been introduced, notably Rubondo Island in
Lake Victoria (P. troglodytes and C. guereza), Ngezi
Forest on Pemba Island and Masingini Forest on Zanzibar
(Unguja) Island (P. kirkii). We made the decision not to
include sites where species had been introduced, although in
some instances this was difficult to prove. For example, it is
not known whether vervets are indigenous to Zanzibar but
there is no irrefutable evidence to the contrary. It is the case
that introductions are different to reintroductions on the
basis that reintroductions are a valid conservation tool.
However, although P. kirkii were introduced to Masingini
(Silkiluwasha, 1981) it is unclear if they were there formerly.
Therefore, we omitted P. troglodytes and C. guereza in
Rubondo and P. kirkii in Ngezi and, as we have no evidence
of their former presence in Masingini, we omitted them at
this site too. For the same reason we retained vervets in the
analysis.
Our study highlighted eight species of particular con-
servation concern in Tanzania: R. kipunji, G. rondoensis,
C. sanjei, P. kirkii, P. gordonorum, Galagoides sp. nov.,
P. troglodytes and G. orinus. We also assigned a threat
category to a number of taxa that had not previously been
designated ( Table 1). However, our main aim was to identify
all key primate sites in Tanzania, to prioritize conservation
intervention and facilitate the role of primates as flagship
or umbrella species (Caro, 2011) for the conservation of
key habitats. The list of Priority Primate Areas identified
differs according to whether species and subspecies, and
diurnal or nocturnal taxa are considered (Supplementary
Tables S2–S3). Top-ranking sites for diurnal taxa are in the
Udzungwa Mountains, Southern Highlands, Zanzibar archi-
pelago, two forested western national parks (Gombe and
Mahale), and the north-western forest of Minziro within the
Guinea-Congo biome. The top-ranking sites for nocturnal
taxa, although also including the Udzungwa Mountains and
Southern Highlands, predominantly include sites in the
coastal forest zone from the East Usambaras in the north to
the south-eastern Lindi forests, as well as the Selous Game
Reserve in south-central Tanzania. There is, therefore, little
overlap in the importance of sites for diurnal and nocturnal
primate species, suggesting that the former cannot be used
as a surrogate for the latter.
Forests are of most relevance for conservation of
primates and are ranked accordingly in our analyses.
This is in contrast to many Important Bird Areas, where
savannah and wetland habitats are often equally important.
However, 44 of the 62 Priority Primate Areas are also Impor-
tant Bird Areas or parts of them. Highest ranking in the
Priority Primate Areas are the species-rich forests of the
Eastern Arc Mountains, the Southern Highlands, includ-
ing Mt Rungwe and Livingstone Mountains, and the
northern ranges incorporating Kilimanjaro, Mt Meru
(Arusha National Park) and Ngorongoro. However, because
of their soils, rainfall and drainage, forested areas are
also important for agriculture and forest products, and
are therefore often zones of the greatest human–primate
conflict.
Based on recent reports of accelerating forest loss, sites
needing particular conservation attention include the forest
reserves of Chome, East Usambaras, Ilole, Mbizi, Milo,
Uzungwa Scarp and Ziwani, and the unprotected forests of
Magombera, Mbuzi, Uzi and Vundwe Islands (Davenport
et al., 2007; Nowak et al., 2009; Nowak & Lee, 2011;Rovero
et al., 2010, 2012). The coastal forests of Pugu and
Kazimzumbwe (two of four forests in the Dar es Salaam
greenbelt; Monga & Gwegime, 2011) are subjected to high
human disturbance because of their proximity to Dar es
Salaam and the city’s demand for charcoal and timber.
However, these forests are potentially manageable as an
urban recreational and educational asset. In general
however, the charcoal industry is growing rapidly and is a
major threat to forests nationwide. Remote locations, such
as the Lindi forests (Perkin et al., 2011 ), are safer from these
pressures but often receive little management attention as a
result.
Of the 62 Priority Primate Areas only five (Ilole,
Madehani, Magombera, Mbuzi and Uzi and Vundwe
Islands) have no official protection status. However, this
gives a misleading impression because few forest reserves in
Tanzania are actively managed in any way. Game Reserve
management usually depends on the zeal of individual
concession holders. Thus not all of the Priority Primate
Areas we propose are equal in terms of conservation
management. Although a national park should be con-
sidered safer than a forest reserve, few protected areas are
managed adequately and adjacent human density can be an
important factor determining conservation success
(Wittemeyer et al., 2008).
The critical subset of Priority Primate Areas (Tables 6–7)
attempts to rationalize through complementarity the full list
of areas and demonstrates how selective targeting could
conserve Tanzania’s primates. The protection of just nine
sites, totalling 8,679 km
2
, would protect all 27 of Tanzania’s
primate species. This subset includes six national parks
(Kilimanjaro, Kitulo, Mahale, Saadani, Udzungwa and
Jozani-Chwaka Bay), one nature reserve (Kilombero) and
Priority Primate Areas in Tanzania 11
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two forest reserves (Minziro and Mgambo). The addition of
just three forest reserves (Rondo, Kilulu Hill and Ngezi) and
two game reserves (Grumeti and Biharamulo) results in a
list of 14 Priority Primate Areas comprising 10,561 km
2
(1.1%) of Tanzania’s total land area, the conservation of
which would ensure the protection of all 43 of Tanzania’s
primate taxa (species and subspecies). We believe therefore
that adoption of the Priority Primate Area system as a
guiding concept could help focus efforts to conserve
primates in Tanzania, and also globally.
Acknowledgements
This study was funded by the Wildlife Conservation Society.
We are grateful to the following people for information and
assistance: Liz Baker, Simon Bearder, Tom Butynski, Tim
Caro, Anthony Collins, Daniela De Luca, Kate Detwiler,
Nike Doggart, Richard Estes, Charles Foley, Trevor Jones,
Yvonne de Jong, Alex Lobora, Jonathan Kingdon, Sophy
Machaga, Noah Mpunga, Arafat Mtui, Alex Piel, the late
Alan Rodgers, Francesco Rovero, Bill Stanley, Fiona St ewart
and Tom Struhsaker.
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Biographical sketches
TIM DAVENPORT’s research interests include biogeography, explora-
tion, herpetology, human impacts, orchidology, primatology and
protected areas. KATE NOWAK studies primate and elephant behav-
iour and the use of refuge habitats in flooded and montane forests.
A
NDREW PERKIN studies the phylogeography, bioacoustics and
taxonomy of galagos with a focus on forested landscapes, and hyrax
and elephant shrew diversity.
Priority Primate Areas in Tanzania 13
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