Arctic, Antarctic, and Alpine Research, Vol. 46, No. 4, 2014, pp. 891–904
Biodiversity and conservation of tropical montane ecosystems in the Gulf of Guinea, West Africa
Drew T. Cronin*†#
Moses B. Libalah‡
Richard A. Bergl§ and
Gail W. Hearn*†
*Department of Biology, Drexel
University, 3245 Chestnut Street, PISB
503, Philadelphia, Pennsylvania 19104,
U.S.A.
†Bioko Biodiversity Protection
Program, Calle Botuku Luba s/n,
Barrio Presidencia, Malabo, Bioko
Norte, Equatorial Guinea
‡Laboratory of Plant Systematics and
Ecology, Higher Teachers’ Training
College, University of Yaoundé I,
Yaoundé, Cameroon
§North Carolina Zoological Park,
4401 Zoo Parkway, Asheboro, North
Carolina 27205, U.S.A.
#Corresponding author:
dtc33@drexel.edu
Abstract
Mount Cameroon (4095 m), the highest peak and only active volcano in West Africa, is
located in the center of the Gulf of Guinea Pleistocene refugium. The associated forests
and highlands along the southern Nigerian-Cameroon border and on the island of Bioko,
known as the Biafran forests and highlands, are important formations of the Cameroon
Volcanic Line owing to their wide elevational range, and on Mount Cameroon, a continuous gradient of unbroken vegetation from sea level to over 4000 m. The montane zones
in the region begin 800 m above sea level forming the critically endangered Mount Cameroon and Bioko Montane Forests ecoregion.
The broad elevational gradient of the region has resulted in high habitat diversity, leading the region to be a center for species endemism and richness across many taxa. Some of
the densest human populations in Africa also occur in this region, putting intense pressure
on the forests and highlands mostly due to overexploitation and habitat loss. The governments of Nigeria, Cameroon, and Equatorial Guinea have designated protected areas in the
region, but coverage is inadequate, especially for the rare montane ecosystems and endemic
taxa. More importantly, protected areas are often not accompanied by effective management
and regulatory enforcement. We recommend improved law enforcement and an expansion
of the protected area network, as well as stronger commitments of institutional, financial,
and technical support from governments and non-governmental organizations, in order to
move conservation in the region in a positive direction. Without significant and immediate
conservation progress, increasing anthropogenic pressure and systemic ineffectiveness of
protected area management represent major concerns for the future of this important area.
DOI: http://dx.doi.org/10.1657/1938-4246-46.4.891
Introduction
The West-African rainforest zone centered between the Cross
and Sanaga Rivers, including Bioko Island, Equatorial Guinea, and
the Cameroon Highlands, has long been recognized for its unique
ecological and biological diversity (Eisentraut, 1973; Barthlott et al.,
1996; Myers et al., 2000; Olson et al., 2001; Oates et al., 2004).
One of the driving factors behind the region’s diversity patterns is
the wide variety of habitats resulting from its extensive highland areas (Fig. 1). The region includes broad interconnected plateaus, like
the Bamenda Highlands, as well as isolated peaks, such as Mount
Cameroon (4095 m) in southwest Cameroon, and Pico Basilé (3011
m) on Bioko Island, the largest insular portion of Equatorial Guinea
(Cable and Cheek, 1998; Oates et al., 2004). Referred to collectively by Bergl et al. (2007) as the Biafran forests and highlands
(BFH), the region has been identified as a center of biodiversity at
both continental (Brooks et al., 2001; Oates et al., 2004) and global
scales (Myers et al., 2000; Olson et al., 2001). The BFH form part of
the West African Forests biodiversity hotspot, and encompass three
ecoregions: the Mount Cameroon-Bioko montane forests, the Cameroon Highlands, and the Cross-Sanaga-Bioko coastal forests (Olson
et al., 2001). High levels of species richness and endemism are represented in the BFH across many taxa, such as primates (Oates, 2011),
amphibians (Lawson, 1993; Schiotz, 1999), birds (Stattersfield et al.,
1998), and vascular plants (Onana and Cheek, 2011). Geographically, the diversity of the BFH is not distributed evenly; patterns of
endemism appear to follow an elevational gradient, with highland
areas harboring the greatest species concentrations (Barthlott et al.,
1996; Oates et al., 2004).
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The biological richness of the BFH is currently under increasing threat from human activities. Although there are no permanent
human settlements within the highest elevation areas of the BFH,
much of the highland zone, which supports the many montane endemic species in the BFH, has no formal protection (Bergl et al.,
2007). Additionally, highland areas are encircled by some of the
highest human population densities in tropical Africa, some of which
exceed 100 inhabitants km–2 (Albrechtsen et al., 2006; CIA, 2013).
These people rely on the forested regions for their health and livelihoods, either directly for their subsistence, or indirectly through
the services provided by those ecosystems (SWPDFW et al., 2005).
High population densities, coupled with a strong rate of population
growth, has led to increased exploitation of remaining forests and
an ever-expanding “human footprint” (Sanderson et al., 2002). This
encroachment has led to the loss of much of the original lowland forest cover and the degradation and fragmentation of many remaining
tracts of forest (Achard et al., 1998; Bergl et al., 2007). Existing protected areas have done reasonably well at protecting habitats more
effectively than alternative land uses (Bruner et al., 2001; Oates et
al., 2004; Struhsaker et al., 2005) thanks, in part, to their relative
isolation and inaccessibility, but habitat loss at the fringes (Achard
et al., 1998; Wittemyer et al., 2008) and hunting within protected
areas are widespread (Fa et al., 2006; Abernethy et al., 2013). Truly
adequate protection will require an expansion of the protected area
network (Table 1) in the BFH and, more importantly, increased efficiency in the enforcement of existing legislation and the management of protected areas within the region.
In this paper, we review the physical history and patterns of
biodiversity and endemism in an effort to assess the current status
DREW T. CRONIN ET AL. / 891
FIGURE 1. Protected areas in the Biafran forests and highlands (BFH). Topography information from the Shuttle-Radar Topography
Mission (SRTM; available from U.S. Geological Survey). Protected area boundaries from IUCN and UNEP (2010).
of threats and conservation progress in the BFH, with an emphasis
on the unique montane ecosystems of the region. We focus particular attention on the “twin peaks” of Bioko Island and Mount Cameroon, due to the authors’ expertise, as well as the peaks’ high elevations, recent shared biogeographic history, and relative isolation
from other highland areas in the BFH (Onana and Cheek, 2011).
We assess the coverage of existing protected areas, as well as major policies that have been established to combat increasing threats
and conserve biodiversity. Finally, we suggest ways in which the
conservation of biodiversity in the region could be improved for
the future.
Geologic and Biogeographic History
The BFH are situated on the margin of the West African and
Congo cratons, where volcanic activity in the Lower Cretaceous (100
Ma) led to the formation of the extensive chain of highlands called the
Cameroon Volcanic Line (CVL) (Tye, 1984). The CVL stretches approximately 1000 km from Lake Chad along a SE-NW axis of continental volcanoes to the volcanic islands of Bioko, Príncipe, São Tomé,
and Annobón, in the Gulf of Guinea (Marzoli et al., 2000; Burke,
892 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH
2001; Tsafack et al., 2009). Its highest formation, Mount Cameroon,
remains the only active volcano in West Africa with seven eruptions
recorded since 1900 (1909, 1922, 1954, 1959, 1982, 1999, and 2000)
(Suh et al., 2003; Tsafack et al., 2009). All other major areas of volcanic activity on the continent are associated with the East African
Rift Valley, over 1800 km away (Cable and Cheek, 1998). The CVL
is unique for being nearly equally divided between the oceanic and
continental lithosphere (Burke, 2001; Tsafack et al., 2009). Mount
Cameroon, on the mainland, and Pico Basilé, on Bioko Island, are
situated on the continental side of the lithospheric boundary, while the
outer Gulf of Guinea islands are oceanic in origin (Jones, 1994; Burke,
2001; Tsafack et al., 2009). The oceanic islands, though not reaching
elevations in excess of 2024 m above sea level (São Tomé), are surrounded by waters approximately 3000 m in depth (Deruelle et al.,
1991). Bioko Island is separated from Cameroon by a 37-km-wide
ocean shelf, which is less than 100 m deep, forming a land bridge
with the African mainland until sea levels rose approximately 10,000
years ago (Jones, 1994; Oates et al., 2004). Patterns of biodiversity
and endemism on Bioko are therefore more similar to those of Mount
Cameroon and mainland Africa than to the outer islands of the Gulf of
Guinea, due to parallels in their recent biogeographic history (Jones,
1994).
TABLE 1
Protected areas of the Biafran forests and highlands.
Area (km2) at:
Protected area
Country
Classification
IUCN PA
Category
Total area
(km2)
0–800
m
800–1700 m
1700–2500
m
>2500
m
Gashaka Gumti NP
Nigeria
National Park
II
5876
3717
2077
82
0
Oban Division Cross
River NP
Nigeria
National Park
II
2687
2661
26
0
0
Douala-Edéa WR
Cameroon
Wildlife Reserve
IV
1681
1681
0
0
0
Korup NP
Cameroon
National Park
II
1295
1288
7
0
0
Kashimbila GR
Nigeria
Game Reserve
IV
1065
972
93
0
0
Mpem et Djim NP
Cameroon
National Park
IV
976
976
0
0
0
Banyang Mbo WS
Cameroon
Wildlife Sanctuary
IV
690
573
116
1
0
Takamanda NP
Cameroon
National Park
II
628
539
89
0
0
Okwangwo Division
Cross River NP
Nigeria
National Park
II
607
557
48
2
0
Mt. Cameroon NP
Cameroon
National Park
II
582
88
254
120
120
Gran Caldera-Southern
Highlands SR
Equatorial
Guinea (Bioko)
Scientific Reserve
Ib
510
289
197
24
0
Pico Basilé NP
Equatorial
Guinea (Bioko)
National Park
II
322
10
224
78
10
Bakossi NP
Cameroon
National Park
II
293
81
207
5
0
Afi Mountain WS
Nigeria
Wildlife Sanctuary
IV
104
84
20
0
0
Santchou WR
Cameroon
Wildlife Reserve
IV
95
44
51
0
0
Kimbi WR
Cameroon
Wildlife Reserve
IV
52
1
51
0
0
Lac Ossa WR
Cameroon
Wildlife Reserve
IV
46
46
0
0
0
Kagwene GS
Cameroon
Gorilla Sanctuary
IV
19
0
10
9
0
17,528
13,607
3470
321
130
Notes: Adapted from Bergl et al. (2007); zonal delineation follows Cable and Cheek (1998). IUCN categories follow Dudley (2008): (Ib) wilderness area—large unmodified or
slightly modified areas, retaining their natural character and influence, without permanent or significant human habitation; (II) national park—large natural or near-natural areas
protecting large-scale ecological processes with characteristic species and ecosystems; (IV) habitat/species management area—areas protecting particular species or habitats, and
management reflects this priority.
Climate
In general, the BFH have a distinctly seasonal climate (tropical equatorial) and rainfall pattern driven by the north-south movement of the Intertropical Convergence Zone (ITCZ) (Oates et al.,
2004). The northward movement of the ITCZ brings heavy rains
from April through October, with a peak between July and September. When the ITCZ is to the south, there is a distinct dry period from November to March that brings dry Harmattan winds
sweeping down from the Sahara (Nosti, 1947; Tchouto et al., 1999;
Oates et al., 2004). The BFH have some of the regions with the
highest mean annual rainfall in Africa, but there is high variation
in local annual rainfall dependent upon topography and proximity
to the coast (Oates et al., 2004). Annual rainfall exceeds 10,000
mm on the southern coast of Bioko and the southwestern foot of
Mount Cameroon, while in the rain shadows, to the north, annual
rainfall is approximately 2000 mm (Nosti, 1947; Tchouto et al.,
1999; Bergl et al., 2007). At least 100 mm of precipitation occurs
each month on the southern coasts of Bioko and Mount Cameroon,
but to the north, in areas like the Obudu Plateau, rainfall may not
exceed 50 mm over a 5 month span (Oates et al., 2004). Due to
the proximity to the equator, the mean annual temperature is about
25 °C with little seasonal variation (Oates et al., 2004). However, elevational gradients can create strong temperature extremes,
ranging from 35 °C at sea level to 4 °C at the summit of Mount
Cameroon (SWPDFW et al., 2005). Persistent high humidity levels
(75%–80%) throughout the year maintain dense cloud cover on the
upper elevations of the southern extent of the region (i.e., Bioko
and Mount Cameroon) (Payton, 1993).
Biodiversity and Endemism
PLEISTOCENE REFUGE
Due to its unique geologic and biogeographic history, the
BFH have been identified as an important Pleistocene refuge area,
which has contributed to its high biodiversity and endemism (Haffer, 1969; Hart et al., 1989; Maley et al., 1990; Oates et al., 2004;
DREW T. CRONIN ET AL. / 893
Anthony et al., 2007). During Pleistocene glaciations, the African
tropics were considerably cooler and drier. Much of the current
lowland closed canopy was open savannah, and the montane zone
extended 1000–1500 m lower than today, occupying significantly
larger areas (Flenley, 1979; Bonnefille et al., 1990; deMenocal,
1995; Gottelli et al., 2004; Assefa et al., 2007). During this period,
the area of montane habitat increased and the distance between
montane habitat patches decreased, which is likely to have facilitated the existence of larger and less isolated populations of species currently restricted to mountains (Moreau, 1963; Assefa et al.,
2007). Roy (1997) suggested that refugia were more impactful on
montane species, leading to rapid divergence of non-continuous
populations, and that montane regions have also acted as centers
of speciation.
FLORA
The Mount Cameroon massif is the only remaining area in
Africa where natural vegetation rises uninterrupted from lowland
forest at sea level to subalpine grassland at the summit (Forboseh et
al., 2011). The southwestern region of the BFH also encompasses
an area of approximately 26,000 km2 of forest that is considered
one of the largest relatively intact contiguous forest blocks in West
Africa (Oates et al., 2004). On a finer scale, however, the structure of the vegetative community of the BFH is highly dependent
on elevation and can differ between sites based on local climate
variation related to features such as latitude, aspect, or proximity
to ocean (Oates et al., 2004). The overall phytogeography of the
region includes formations dominated by Guineo-Congolian and
Lower Guinea rain forest species, with Afromontane elements at
higher elevations, and can be broadly categorized into strata according to elevation (Table 2) (Fa, 2000).
Plant species diversity in the BFH is the highest in tropical
Africa (Barthlott et al., 1996), owing largely to its varied habitat
mosaic. Mount Cameroon, for example, is an especially speciose
center of plant diversity, with a total of 2435 species of vascular
plants, relative to 1693 species in nearby lowland Korup National
Park, and 1105 species (angiosperms only; Figueiredo, 1994) on
Bioko Island (Cable and Cheek, 1998; Onana and Cheek, 2011).
There is high affinity between the plant species of Bioko and western Cameroon, which suggests that Bioko is floristically part of the
mainland (Exell, 1973). There are no strict endemic plants at the
upper extent of Mount Cameroon (3500–4095 m); however, Cable
and Cheek (1998) listed a total of 49 total endemics for the massif,
of which 20 are montane species (11: 800–1800 m; 5: 1800–2100
m; 4: 2100–3500 m). There are four montane grassland endemics, of which two (Silene biafrae [Caryophyllaceae], Hypseochloa
cameroonensis [Gramineae]) are listed as vulnerable and two (Bulbostylis densa var. cameroonensis [Cyperaceae], and Habenaria
obovata [Orchidaceae]) are recognized as endangered (Onana and
Cheek, 2011; IUCN, 2013). Relative to Mount Cameroon, whose
49 endemics constitute 2.01% of its overall species number, Bioko
Island has at least 40 endemic species, giving it a higher relative
level of endemism (3.62%) (Figueiredo, 1994).
FAUNA
The BFH are a hotspot for faunal species richness and endemism across taxonomic groups (Myers et al., 2000; Brooks et
al., 2001). As a result, the Cameroon Highlands are considered
one of the top five conservation priorities in Africa for terrestrial
vertebrates (Brooks et al., 2001), the Mount Cameroon and Bioko
montane forests ecoregion is among the most important for the
conservation of forest-dependent bird species (Buchanan et al.,
TABLE 2
Generalized forest type strata of the Biafran forests and highlands (BFH) with corresponding altitudinal range, coverage extent, and
proportion occurring within protected area boundaries, and characteristic species.
Forest type1
Altitudinal
range (m)1
Extent
covered (m)1
Proportion protected
(%)
Lowland forest
0-800
123,731
11.00
Ficus spp. are dominant; Chlorophora excelsa; Chrysophyllum
africanum; Strombosia scheffleri; Symphonia globulifera;
Caesalpinioideae (Leguminosae)1,2,3
Submontane forest
800-1700
53,407
6.50
Alangium chinense; Cyathea camerooniana; Cyathea
manniana; Polyscias fulva; Psydrax dunlapii; Oncoba
lophocarpa; Oncoba ovalis; Xylopia africana; large tracts
of monocarpic Acanthaceae (e.g. Mimulopsis solmsii);
Afromomum spp. and Maranthaceae prominent in understory1
Upper montane forest
1700-2500
4,034
7.96
Schefflera abyssinica; Schefflera mannii; Prunus africana;
Xymalos monospora; Hypericum revolutum; Clausena
anisata; Nuxia congesta; Afromomum spp. and Maranthaceae
prominent in understory1,4
Montane scrub
2500-2800
99
48.72
Erica mannii, Erica tenuicaulis, Agarista salicifolia;
Hypericum roeperianum, Hypericum lanceolatum; Gnidia
glauca, Maesa lanceolata, and Myrica arborea1,3,5
Subalpine grassland
>2800
89
92.25
Veronica mannii; Pentaschistis mannii; absence of Loudetia
simplex; presence of temperate plant genera including
Helichrysum spp., Geranium spp., Clematis spp., Senecio spp.,
and Solanum spp.1,3
Common species1
Forest types, altitudinal ranges, and common species are approximated from Cable and Cheek (1998). Information on the extent of coverage approximated from the Shuttle-Radar
Topography Mission (SRTM; available from U.S. Geological Survey). Protected area boundaries from IUCN and UNEP (2010).
1
Cable and Cheek (1998); 2Richards (1963); 3Fa (2000); 4Richards (1996); 5Leuschner (1996).
894 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH
2011), and Bioko Island has been ranked as the single most important place in Africa for the conservation of primate diversity
(Oates, 1996). Biodiversity and endemism patterns within the BFH
vary widely between taxa but seem linked to terrain and dispersal
ability. For example, primate endemism is highest in the lowlands,
where rivers appear to be a major dispersal barrier. Birds, on the
other hand, are not restricted by rivers, but do exhibit high levels
of montane endemism, largely due to the relative isolation between
montane areas in the BFH (e.g., ~50 km between Mount Cameroon
and Pico Basilé) and from any similar region in Africa (Oates et
al., 2004). A number of endemic taxa are present in current protected areas, although the variety of endemism patterns across taxa
has led to a disconnect between faunal distributions and protected
areas. The majority of taxa endemic to the BFH are montane, yet
insufficient highland area is formally protected (Bergl et al., 2007).
The following faunal overview follows Bergl et al. (2007), focusing primarily on primates, birds, and amphibians, as these taxa are
better studied and adequate data were available.
Mammalian species, and especially primates, are particularly
well represented in the region (see Oates et al. [2004] for a descriptive list). A total of 32 primate taxa are distributed across the BFH,
including 13 endemics, of which 8 are endangered and 2 are critically
endangered (Oates, 2011; IUCN, 2013). Numerous primate species
inhabit highland areas throughout the BFH, but although Preuss’s
monkey (Allochrocebus preussi) is primarily associated with montane
forest, there are no strict montane endemics (Oates, 2011). Patterns of
montane endemism in mammals in the region are perhaps best represented in the distribution and elevational range of endemic rodents
across highland areas in the BFH (Table 3). For example, seven species across three genera, Crocidura, Myosorex, and Sylvisorex, comprise the endemic Soricidae taxa. Each of these species exhibits a distribution confined to montane habitats in either a single highland area,
or small series of highlands (Fig. 2) (IUCN, 2013).
The BFH have the highest bird species richness in west and
central Africa due to the overlap of Upper and Lower Guinea
species and the spectrum of habitats afforded by the elevational
range and topography of the highlands (Smith et al., 2000; Oates
et al., 2004). Furthermore, localized estimates of species richness
(Bioko and western Cameroon: Eisentraut [1973]; Korup: Green
and Rodewald [1996]) are believed to be an underestimate of the
total number of bird species in the region (514 species; Myers et
al., 2000; Oates et al., 2004). Avian endemism is high, but there
FIGURE 2. Distribution
of montane endemic rodents
(Soricidae) in the BFH.
Distribution data from
IUCN (2013). Protected
area boundaries from IUCN
and UNEP (2010).
DREW T. CRONIN ET AL. / 895
TABLE 3
Endemic montane rodents of the Biafran forests and highlands, their altitudinal range, and IUCN Red List category.
Binomial
Altitudinal Range
(m)
Common name
IUCN
Category
Muridae
Hybomys badius
Eisentraut’s Striped Mouse
> 800
EN
Hybomys basilii
Father Basilio’s Striped Mouse
450–2000
EN
Hylomyscus grandis
Mt. Oku Hylomyscus
2100
CR
Lamottemys okouensis
Mt. Oku Rat
2100–2900
EN
Lemniscomys mittendorfi
Mittendorf’s Striped Grass Mouse
2100–2300
VU
Lophuromys dieterleni
Dieterlen’s Brush-furred Mouse
2100
EN
Lophuromys eisentrauti
Eisentraut’s Brush-furred Mouse
2550
EN
Lophuromys roseveari
Roseveari’s Brush-furred Mouse
1000–3100
LC
Otomys burtoni
Burton’s Vlei Rat
2000–4000
EN
Otomys occidentalis
Western Vlei Rat
1900–3000
VU
Praomys hartwigi
Hartwig’s Soft-furred Mouse
2700–2900
EN
Praomys morio
Cameroon Soft-furred Mouse
1100–2135
EN
Praomys obscurus
Gotel Mountain Soft-furred Mouse
1600–2400
EN
Cooper’s Mountain Squirrel
~800–2500
DD
Crocidura eisentrauti
Eisentraut’s Shrew
2000–3000
VU
Crocidura picea
Cameroonian Shrew
1200–1800
EN
Myosorex eisentrauti
Eisentraut’s Mouse Shrew
>2000
CR
Myosorex okuensis
Oku Mouse Shrew
1800–2300
EN
Myosorex rumpii
Rumpi Mouse Shrew
1100
EN
Sylvisorex camerunensis
Cameroonian Forest Shrew
2000–2300
VU
Sylvisorex morio
Mt. Cameroon Forest Shrew
>1200
EN
Sciuridae
Paraxerus cooperi
Soricidae
Notes: Species occurrence from Kingdon, 1997; Maisels et al., 2001; Oates et al., 2004; Amori et al., 2008. Approximate altitudinal range from IUCN, 2012. Species
recorded from a single elevation are listed as single values. IUCN Red List categories: critically endangered (CR), endangered (EN), vulnerable (VU), least concern (LC),
data deficient (DD).
is little consistency in distribution patterns among endemic taxa,
apart from exhibiting a preference for montane forests and grasslands (Bergl et al., 2007). Only three species are recorded from a
single montane site: the Mount Cameroon francolin (Francolinus
camerunensis) and the Mount Cameroon speirops (Speirops melanocephalus) from Mount Cameroon, and the Fernando Po speirops (Speirops brunneus) from Pico Basilé (Pérez del Val et al.,
1994; Oates et al., 2004; IUCN, 2013). Of the 26 regional endemics, 58% are currently threatened (6: endangered; 4: vulnerable; 5:
near threatened) (IUCN, 2013).
Myers et al. (2000) estimated 139 reptile species and 116
amphibian species occur in the West African forests hot spot.
Eisentraut (1973) listed 52 reptile and 32 amphibian species
from Bioko, while Lawson (1993) listed 83 reptile and 90 amphibian species in Korup National Park. Similar to the total
number of bird species in the region, it is suggested that the
overall species richness of herpetofauna in the region is currently underestimated and may actually be considerably higher
(Oates et al., 2004). Amphibians are relatively better studied in
896 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH
the region and exhibit higher estimates of endemism (77%) than
reptiles (33%) (Myers et al., 2000). In contrast to the respective lowland- and montane-centered distribution of the primates
and birds, Gartshore (1984) and Bergl et al. (2007) described
a vertically stratified distribution of endemic amphibians, with
distinct lowland, lower montane, and upper montane species.
Of the 53 species endemic to the region, 39 (73.6%) species are
recorded only above 800 m. Twelve (30.1%) of these species
are restricted to an altitudinal range of 800–1600 m, 16 (41.0%)
are found only over 1200 m, and among these, 10 (25.6%) are
found only at altitudes greater than 1600 m (Bergl et al., 2007;
Zimkus, 2009; Blackburn, 2010).
Protected Areas
The BFH contains 18 strict protected areas, comprising
three International Union for Conservation of Nature (IUCN)
categories (Ib, Scientific Reserve; II, National Park; IV, Wild-
life Sanctuary), and encompassing a total area of over 17,500
km 2 (Table 1). Based on the breakdown of vegetation strata in
Cable and Cheek (1998) (Table 2), approximately 3921 km2
(22%) are at an elevation above 800 m, 451 km2 (2.6%) are
above 1700 m, and 446 km 2 (0.74%) are above 2500 m. It is
also noteworthy that all land above 1600 m on Bioko Island is
in protected areas. Gashaka Gumti National Park in Nigeria
encompasses a majority (55%) of the total protected highland
area; however, the park lies outside the Guineo-Congolian
moist forest zone, with only small patches of montane forest
and limited habitat for endemic montane species (Bergl et al.,
2007). Overall, it is estimated that only 6.0% of approximately
65,000 km2 of highland ecosystems above 800 m in the region
have any formal protection (Bergl et al., 2007).
Effective conservation within established protected areas
is uncommon. Existing protected areas, often by nature of their
terrain, have been relatively successful in protecting large tracts
of habitat (Bruner et al., 2001; Oates et al., 2004; Struhsaker et
al., 2005); however, they are under intense threat from burning,
agriculture, livestock grazing, and, most especially, the hunting
of larger vertebrates, such as anthropoid primates and ungulates
(Maisels et al., 2001; Chapman et al., 2004; Oates et al., 2004;
Fa et al., 2006; Linder and Oates, 2011; Abernethy et al., 2013;
Cronin, 2013; Cronin et al., 2013). Moreover, many of the region’s
protected areas lack both clarity in their legal boundaries and any
effective management plan (Oates et al., 2004). Indeed, many exist solely as “paper parks,” where conservation and management
activities are limited or nonexistent (Blom et al., 2004; Oates et
al., 2004; Bergl et al., 2007; Cronin et al., 2010). On Bioko, for
example, there is no management plan in place for the Gran Caldera–Southern Highlands Scientific Reserve (GCSH), the protected
area with the highest IUCN designation (Ib; Table 1) in the BFH.
The GCSH boundary remains unmarked, and, in addition to the
absence of park rangers or management staff, the few military personnel responsible for law enforcement within the reserve regularly hunt primates within its boundaries (Cronin, 2013). On Mount
Cameroon, despite the creation of a national park in 2010 (Forboseh et al., 2011), a management plan has yet to be implemented,
much of the boundary remains unmarked, and regular exploitation
from surrounding populations remains common. Ultimately, active
noncompliance and the absence of effective management occur
throughout the BFH, essentially nullifying much of the value of
gazetting a protected area (Bergl et al., 2007).
Human Population
The BFH support some of the most densely populated areas
on the continent. Nigeria is the second most densely populated
country in Africa (184 people km–2), with densities upward of 500
people km–2 in some southeastern areas along the Cameroon-Nigeria border (Oates et al., 2004; CIA, 2013). Cameroon is less densely populated (42 people km–2), but the Bamenda Highlands, which
lie entirely within the study region, are one of the most densely
populated areas in the country (Oates et al., 2004; CIA, 2013). For
instance, Mount Cameroon, the most unique formation in the CVL,
is estimated to support 300,000 individuals (SWPDFW et al.,
2005). Human settlements consisting of high-density urban areas
and smaller villages form a ring with little remaining forest cover
around its base up to 1500 m in places (Fotso et al., 2001; SWPDFW et al., 2005). The population of Bioko Island is estimated at
roughly 180,000 people, with approximately 137,000 people living in and around the northern capital city of Malabo (CIA, 2013;
Cronin, 2013). The remainder of the island’s population lives in
villages and towns encircling Pico Basilé at low elevations and on
the northern flanks of the GCSH, with population densities less
than 10 people km–2 in the south (Albrechtsen et al., 2006).
Threats
The threat to biological diversity is high in West Africa, relative to other places in sub-Saharan Africa, as a result of high human
population density and growth rate, as well as a high rate of habitat
loss (Brashares et al., 2001; Wittemyer et al., 2008). Wittemyer et
al. (2008) suggested human settlements around protected areas are
strong predictors of illegal timber and mineral extraction, bushmeat hunting, fire frequency, and species extinctions. Exacerbating
the situation is that protected areas seem to attract human settlement, as rates of population growth surrounding protected areas are
nearly double that of average rural growth rates (Wittemyer et al.,
2008). The associated increase in anthropogenic activities, especially deforestation and bushmeat hunting, in the BFH has had progressively more deleterious effects on the biodiversity and fragile
ecosystems of the region (Achard et al., 1998; Oates et al., 2004).
DEFORESTATION
Although there has been considerable deforestation and forest degradation in the BFH associated with development and the
expansion of subsistence activities, such as agriculture, energy
(e.g., fuelwood), and timber (Charlotte, 2010; de Wasseige et al.,
2012; Megevand et al., 2013), Africa has contributed considerably
less overall (5.4%) to the global loss of humid tropical forests relative to Asia and the Neotropics (Hansen et al., 2008). The annual
net deforestation rate in the Congo Basin has accelerated recently,
however, with losses corresponding to about 0.17%, or approximately 300,000 km2, each year (de Wasseige et al., 2012). Deforestation estimates for Cameroon suggest the loss of approximately
800–1000 km2 per year (Alpert, 1993; Wolfe et al., 2005), with the
coastal region suffering the most intensive exploitation (Laporte
et al., 2007; de Wasseige et al., 2012). Remote sensing has also
indicated that Cameroon and Equatorial Guinea had the greatest
densities of logging roads (0.09 km km–2) and the greatest amount
of forest disturbance (15%) in Central Africa, while the Mount
Cameroon and Bioko montane forests had the highest percentage
of mean forest loss from 2000–2005 (2.40%), out of the 20 ecoregions most important for the conservation of forest-dependent bird
species (Buchanan et al., 2011). High human densities and continued human immigration to the area have driven this trend and led
to the clearance of much of the natural vegetation for both subsistence and commercial agricultural use, while the majority of lowland forests have been cleared for industrial plantations, such as
oil palm (Elaeis guineensis) (Forboseh et al., 2011; Linder, 2013).
Deforestation was once widespread on Bioko, as nearly 60% of its
lowland forests were cleared for cocoa and other tropical crops;
however, nearly half of the converted land has since been abandoned for agricultural use and has been reclaimed by scrub and
secondary forest (Butynski and Koster, 1994).
The higher elevations of both Mount Cameroon and Bioko
remain largely intact due to their low potential value for exploitation and their relatively inaccessible rugged terrain (Butynski and
Koster, 1994; Fotso et al., 2001; Oates et al., 2004). As a result,
no major human activities or settlements occur above 2000 m.
On Mount Cameroon, paved roads reach Buea (870 m), but go no
DREW T. CRONIN ET AL. / 897
further. On Bioko, Moeri (720 m) is the highest permanent settlement on Pico Basilé, however, a guarded road provides access to a
meteorological and telecommunications facility and its associated
military installation at the summit. The village of Moka (1400 m)
is the highest overall on Bioko, situated at the northern border of
the GCSH.
BUSHMEAT HUNTING
Bushmeat hunting is extensive and unsustainable throughout
the BFH (Fa et al., 2000, 2006; Albrechtsen et al., 2007; Morra et al.,
2009; Linder and Oates, 2011; Cronin, 2013; Cronin et al., 2013),
threatening many large vertebrates with extinction, especially primates (IUCN, 2013). It is a highly commercialized activity, fueled
by human population growth and increased per capita wealth in
urban centers, modernized hunting techniques, and increased accessibility to remote areas of forest (Robinson and Bennett, 2000;
Albrechtsen et al., 2007). The magnitude of faunal exploitation
is great; over 197,000 carcasses were counted from Bioko from
1997–2010 (Cronin, 2013), while Fa et al. (2006) recorded over
42,000 kg of bushmeat in Cross-Sanaga region of the mainland in
a six-month study period alone. Hunting has a negative impact on
the diversity and densities of large-bodied vertebrates and can lead
to adverse and cascading effects on ecosystem functioning (Redford, 1992; Chapman and Onderdonk, 1998; Wang et al., 2007;
Vanthomme et al., 2010; Abernethy et al., 2013). Although much
of the region is classified as protected (e.g., ~42% of Bioko Island),
legislation aimed at restricting hunting has failed, due to a lack
of management and to ineffective or absent enforcement regimes
(Oates et al., 2004; Struhsaker et al., 2005; Bergl et al., 2007).
CLIMATE CHANGE
It is projected that climate change will most severely affect
the African continent (IPCC, 2014b), particularly in the central African region of the BFH (Penlap et al., 2004; James et al., 2013).
Warming projections suggest the rise in mean annual temperature
is likely to exceed 2 °C across large swaths of the continent under
medium scenarios, and its entirety under high-emission scenarios
(IPCC, 2014b). Highland areas, such as the BFH, will be especially
affected, as warming is expected to be more intense relative to lowlands (Pounds et al., 1999), and rainfall patterns are predicted to
change dramatically (IPCC, 2014b). Indeed, montane ecosystems
throughout Africa are already responding to climate change (Chen
et al., 2009; Allen et al., 2010; Eggermont et al., 2010; Chen et al.,
2011; Laurance et al., 2011; Willis et al., 2013; IPCC, 2014b).
Global modeling studies have predicted that over 30% of
plant and animal species will be threatened with extinction given
a rise in mean annual temperature in excess of 1.5 °C (Thomas et
al., 2004). These extinctions will be disproportionately attributed
to tropical areas (Thomas et al., 2004; Colwell et al., 2008), due to
a number of factors including species richness and high endemism
(Colwell et al., 2008; Raxworthy et al., 2008). Because the effects
of climate change are predicted to be amplified in highland areas
(Pounds et al., 1999; IPCC, 2014b), tropical montane zones will
likely be particularly affected (Colwell et al., 2008; Ohlemüller et
al., 2008; Raxworthy et al., 2008). Climate models for the tropics
suggest that the coolest climatic zones at the upper elevations will
be lost (IPCC, 2014a), and that there will be a shift of remaining
vegetation strata upslope threatening corresponding species and
montane endemics with extinction (Still et al., 1999; Beniston,
2000; Thomas et al., 2004; Raxworthy et al., 2008; Sekercioglu
898 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH
et al., 2008; Chen et al., 2009). Montane endemics will be faced
with substantial range contractions, increasing prevalence of
climate-driven infectious disease (Pounds et al., 2006), and even
“‘mountain top”‘ extinctions (Pounds et al., 1999; Colwell et al.,
2008), resulting from limited dispersal capabilities (Laurance et
al., 2011), narrow ranges (Ohlemüller et al., 2008), and restricted
physiological tolerances (Beniston, 2000; Schloss et al., 2012).
For example, species like Hartwig’s soft-furred mouse (Praomys
hartwigi), currently known from a highly restricted elevational
range (2700–2900 m) just below the summit of Mount Oku (3011
m), may have difficulty adapting to rapid environmental change.
Anthropogenic impacts are expected to exacerbate the effects
of climatic change (Bush, 2002; Colwell et al., 2008), as bushmeat
hunting is interfering with forest regeneration and seed dispersal
(Wilkie et al., 2011; Abernethy et al., 2013), and rapid habitat loss
and fragmentation are disrupting dispersal capabilities (Achard
et al., 1998; Bergl et al., 2007; Laporte et al., 2007; Bergl et al.,
2008; Laurance et al., 2009). Given current levels of habitat loss
and anthropogenic pressure, the higher elevations of the BFH are
increasingly becoming “sky islands,” acting as refuge (Pounds et
al., 1999; Chen et al., 2009) from increasing encroachment from
the lowlands, but isolated from other highland areas (Butynski et
al., 1997; Newmark, 2008).
OTHER THREATS
Additional threats to BFH include fires and volcanic eruptions, as well as the unregulated collection of non-timber forest
products, including honey, wild vegetables, and medicinal plants.
The high-elevation vegetation communities of the BFH are prone to
damage by fire and exhibit slower growth rates and natural regeneration than other regions, making the effects of even short-lived
fire events long-lasting (Charlotte, 2010; Forboseh et al., 2011).
Fires in the montane zone can be of natural (e.g., lighting, volcanic eruption) or anthropogenic origin (e.g., hunters flushing out
game, honey collectors flushing out bees) (Forboseh et al., 2011).
Anthropogenic fire events are readily observable and often grow
swiftly out of control (Cronin; Libalah, personal observation). The
collection of non-timber forest products, such as African jointfir
(Gnetum africanum), the fruits of Afromomum spp., bush mango
(Irvingia gabonensis), and African whitewood (Enantia chlorantha), is also common throughout the BFH (Charlotte, 2010). Even
so, the exploitation of the montane scrub and subalpine grasslands
in the region has been primarily restricted to hunting of game. Recent findings by Zofou et al. (2011), however, justified the use of
stem bark from Hypericum laceolatum (Hypericaceae), found in
the upper montane zone of the BFH (Table 2), for the treatment
of malaria, and suggest that it will likely yield new anti-malarial
drug candidates. Given the gravity of malaria infection worldwide,
further positive results may lead to local overexploitation similar
to that of another montane species, the red stinkwood (Prunus africana), whose bark is used to treat prostate hyperplasia (Ingram and
Nsawir, 2007; Charlotte, 2010).
Environmental Legislation
Ineffective protected area management is rampant in the BFH.
Widespread illegal exploitation of the resources within protected
areas results from a myriad of factors, such as unclear borders, lack
of enforcement, limited institutional capacity, and inadequate financial resources (Oates et al., 2004; Njuh Fuo and Memuna Semi,
2011). Additionally, those tasked with legislation and enforcement
are often either underpaid or involved in the exploitation—by consuming the resource in question, turning a blind eye, accepting
bribes, actively hunting, or falsifying official documents regulating resource use (Nguiffo and Talla, 2010; Peh and Drori, 2010;
Cronin, 2013). Unfortunately, the inability to effectively impose
legislation appears common in Africa, despite well-intentioned efforts from numerous individuals, non-governmental organizations
(NGOs), conservation departments, and governments (Peh and
Drori 2010).
The environmental legislation of Equatorial Guinea provides clear insight into the underlying systemic mismanagement
of the region. Equatorial Guinea has passed four major laws on
the environment (Republic of Equatorial Guinea, 1988, 2000,
2003, 2007). Laws No. 8/1988 (Hunting, Wildlife, and Protected
Areas) and No. 4/2000 (Protected Areas) were both superseded
by No. 7/2003 (Environmental Regulation), which tasked a new
government agency, INCOMA/FONAMA, with the responsibility of managing protected areas. To date, INCOMA/FONAMA
does not exist and there is no enforcement of the law’s provisions. Articles (34, 36, 37, and 46) of Law No. 7/2003 also cover
the same tenets as Decree No. 72/2007, which bans the hunting,
sale, and consumption of primates. Furthermore, both Pico Basilé
National Park and the GCSH lack management plans, an urgent
conservation concern (Cronin et al., 2010). Given the unclear nature of environmental law, jurisdiction, and protected area management in Equatorial Guinea, it is not surprising that there has
been little conservation progress via legislation.
The main legal framework for environmental management
in Cameroon is Law No. 96-12 of 5 August 1996 (Republic of
Cameroon, 1996); however, there are a number of policies that
regulate specific environmental sectors. The “‘wildlife code”‘
was established through Law 94-01 of 19 January 1994 (Republic
of Cameroon, 1994), which provides a legal code for the use of
forests, wildlife, and fisheries, and Decree 95-466-PM of 20 July
1995 (Republic of Cameroon, 1995), which specifies the conditions for the implementation of Law 94-01 (Nguiffo and Talla,
2010; Njuh Fuo and Memuna Semi, 2011). Similar to the example given above for Equatorial Guinea, the wildlife code also
suffers from a number of shortcomings. For instance, effective
implementation of the wildlife code is dependent on “‘enabling
decrees”‘ (Republic of Cameroon, 1994), a number of which
have not been enacted, and can sometimes take years to be put
into effect (Njuh Fuo and Memuna Semi, 2011). The wildlife
code also mandates that logging companies must develop forest
management plans for each of their forest parcels and submit it to
the Ministry of Forests and Fauna (MINFOF) for approval within
three years of allocation (Republic of Cameroon, 1994), but many
of the approved management plans do not comply with minimum
legal prescriptions (Cerutti et al., 2008), and critics argue that
the delegation of forest surveys to logging companies has sacrificed the environment for economic considerations (Njuh Fuo
and Memuna Semi, 2011). Another central tenet of the wildlife
code obliges the government to classify animal species into three
classes, according to their level of protection (Republic of Cameroon, 1994), and requires that the classification is updated every
five years (Republic of Cameroon, 1995). Despite the requirement, the government has not regularly updated the classification,
which diminishes the currency and reliability of data available to
policy-makers and management professionals (Nguiffo and Talla,
2010). Furthermore, despite legislative classification as ‘Class
A’ species, that may on no occasion be killed, illegal hunting of
wildlife, such as the endangered chimpanzee (Pan troglodytes)
and critically endangered gorilla (Gorilla gorilla), is extensive
(Fa et al., 2006; Bergl et al., 2011; Djeukam et al., 2012).
Recommendations
The threats facing the BFH are multifaceted and may require
localized strategies to best manage resources. Hunting mitigation
strategies, for example, should vary given the primarily commercial nature of the trade on Bioko relative to Cameroon, where a
greater proportion is subsistence based. However, across the BFH
there is a commonality of requirement for improved law enforcement, and strong commitments to environmental protection from
governments and NGOs, by way of institutional, financial, and
technical support (Struhsaker et al., 2005; Njuh Fuo and Memuna
Semi, 2011; Cronin, 2013).
Increased effectiveness of law enforcement is of paramount
importance to the conservation of the BFH (Oates et al., 2004;
Struhsaker et al., 2005; Bergl et al., 2007; Bennett, 2011; Wilkie
et al., 2011; Tranquilli et al., 2012), to which the most practical
short-term solution is the implementation of forest guards (Bennett, 2011). Forest guards have been a successful strategy that has
been linked to reductions in hunting and improved effectiveness
of protected areas (Bruner et al., 2001; Rowcliffe et al., 2004; de
Merode and Cowlishaw, 2006; Hilborn et al., 2006; Bennett, 2011;
Campbell et al., 2011; Tranquilli et al., 2012). An expansion of
the protected area network, as well as increasing the size of existing reserves, will also be essential to the conservation of the BFH.
Many protected areas in the region are too small and are suffering
from levels of exploitation that are too high to sustain populations
of many species (Brashares et al., 2001; Struhsaker et al., 2005).
Increasing the size of protected areas will reduce the area to edge
ratio, as well as hunter accessibility to the core of the reserve. Furthermore, an expansion of the protected area network to provide
coverage for the inadequately protected highland ecosystems and
endemic taxa in the BFH would greatly improve conservation
overall in the region (Bergl et al., 2007). New protected areas, or
an expansion of existing protected areas, like the proposed corridor linking montane areas of GCSH and PNBP on Bioko (UNDPGEF, 2010), will be important, but effective management and law
enforcement in existing protected areas is the most critical factor
for the conservation of biodiversity in the BFH (Struhsaker et al.,
2005; Bergl et al., 2007; Cronin et al., 2010). The primate hunting
ban on Bioko, for instance, includes prohibitive fines that, if enforced, would threaten the entire estimated annual hunting income
for hunters and make it uneconomical for both suppliers and consumers alike to persist (Fa et al., 2000).
The cost of biodiversity conservation is minimal relative to
the value of the ecosystems being protected (James et al., 1999),
with estimates suggesting that conservation in protected areas
could be effectively achieved for just 1% of the annual value of
natural ecosystems (Pimentel et al., 1997). This is particularly true
in the BFH, where the total costs of biodiversity conservation are
minuscule in comparison to estimates of profits from environmental exploitation (e.g., industrial logging accounts for 11% [~$3 billion] of the GDP of Cameroon) (Huarez et al., 2013; World Bank,
2014a). A 2005 assessment (Struhsaker et al., 2005) identified that
most African rain forest protected areas were underfunded, and at
least 75% lacked a secure long-term funding source despite significant involvement from international donors. This represents
a long-term concern, but also identifies a glaring problem with
protected area funding in the BFH, lending further support to the
lack of a credible commitment from regional governments to en-
DREW T. CRONIN ET AL. / 899
vironmental protection. The cost of operating a protected area in
African rain forest was between $23 and $208 km–2 in 2005, and
even doubling those estimates to $400 km–2 would still have left
the costs significantly lower than protected areas in developed nations (James et al., 1999; Struhsaker et al., 2005). Adjusted for
inflation, $400 km–2 would be approximately $490 km–2 in 2014,
which results in a projected operating cost of just over $8.5 million
for all identified protected areas in the BFH (Table 1), less than
1% of the gross profits from timber in Cameroon. On Bioko, the
estimated annual cost of operating its two protected areas would
be just $408,000, only 0.003% of the overall GDP of Equatorial
Guinea (World Bank, 2014b). There are, of course, myriad factors that govern protected area funding, and the values presented
here are simply an estimate. However, despite Cameroon’s leading
role in Congo Basin forestry legislation (Cerutti et al., 2008) and
the designation of environmental conservation as one of Equatorial
Guinea’s ‘Five Pillars’ of reform (Qorvis, 2010), these estimates
are illustrative of the lack of funding allocated to protected areas in
the BFH and the relatively low cost at which environmental protection in the region could operate efficiently. True commitment to
conservation in the BFH will ultimately require greater financial
investment from regional governments. Moreover, future funding
structures need to be both secure and long-term, such as trust funds
or endowments, where the annual return on investment will continue to supply funding for the protected area over time.
Many of the flaws of environmental conservation and management in the BFH ultimately stem from governments that are
lacking in political will (Smith et al., 2003; Cerutti et al., 2008;
Njuh Fuo and Memuna Semi, 2011; de Wasseige et al., 2012), and
conservation departments that have little political clout. The empowerment of conservation departments will be critical in order
for them to more effectively combat environmental offenses by
citizens and other sectors (Smith et al., 2003). Situations, such as
false CITES (Convention on the International Trade in Endangered
Species) certificates for the export of 1200 parrots from Cameroon
(Nguiffo and Talla, 2010), or illegal permits signed by senior military officials for the poaching of marine turtles on Bioko (Cronin,
personal observation), can only truly be combated if the perpetrators cannot act with impunity, and can be prosecuted by conservation departments to the fullest extent of the law. Moreover, institutional corruption, a problem throughout the BFH, can detract from
the conservation progress and lower the effective funding available
for conservation initiatives (Smith et al., 2003; Struhsaker et al.,
2005). Efforts have been made to address corruption in the BFH by
organizations, such as the Last Great Ape Organization (LAGA),
which have had success in lobbying for the enforcement of environmental laws in Cameroon; however, fixing the institutionalized corruption common in the BFH will require a governmental
overhaul, as well as the political will and leadership to see such a
divisive undertaking through (Peh and Drori, 2010).
NGOs also play an essential role in environmental conservation and law enforcement in the BFH, and going forward, their
role will only be greater. Organizations, such as LAGA, Wildlife
Conservation Society (WCS), and the World-Wide Fund for Nature
(WWF), have been instrumental in helping to bring environmental
offenders to justice (Njuh Fuo and Memuna Semi, 2011), and technical and/or financial NGO support has been strongly linked to the
creation and some degree of success of protected areas (Struhsaker
et al., 2005). Meanwhile, many smaller NGO’s, like the Central
African Biodiversity Alliance (CABA) in Cameroon and the Bioko
Biodiversity Protection Program (BBPP) on Bioko, have been successful by partnering with local institutions and promoting conservation through education and research. Expanding long-term NGO
900 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH
involvement and partnerships through further proliferation of research initiatives in the BFH appears to be one viable path toward
immediate on-the-ground conservation success. Studies suggest
that effective conservation can be achieved through the establishment of a research presence (Campbell et al., 2011; N’Goran et
al., 2012). Successful projects, like San Diego Zoo Global’s Ebo
Forest Research Project and WCS’s gorilla monitoring work at
Kagwene Gorilla Sanctuary, demonstrate the value of research for
conservation in the BFH. Expanded research initiatives in the BFH
could also generate data critical for improving conservation management and future planning (e.g., N’Goran et al., 2012), which are
lacking for many protected areas (Struhsaker et al., 2005).
While an expansion of NGO-led research could have significant
conservation impacts, it is clear that the extent of threat in the BFH
will also require considerable law enforcement intervention in order
to secure the region’s biodiversity. Increasingly, evidence suggests that
effective conservation in tropical Africa must be tied to effective protection programs (e.g., Holmern et al., 2007; Fischer, 2008; Jachmann,
2008; Tranquilli et al., 2012). Lack of capacity and political will on
the part of governments in the region has provided opportunities for
NGOs to play significant roles in the support and management of protected areas in the BFH. Both WCS and WWF are engaged at various
levels with conservation law enforcement in Cameroon, and in Nigeria, WCS directly manages ranger programs, in partnership with government agencies and community groups, at two sites. Public-private
partnerships such as these will likely be the best way to control threats
to biodiversity and prevent local extinctions in the immediate term.
More broadly focused institutions, such as the Central African
Forests Commission (COMIFAC) and Congo Basin Forest Partnership (CBFP), have also leveraged the collective expertise of their
numerous stakeholders in order to promote regional scientific exchange and collaboration on conservation action. Moving forward,
multi-stakeholder, regional planning of unified and comprehensive
conservation strategies will be critical to the future of the BFH, as
mitigation of transboundary issues, such as wildlife trade, biodiversity loss, and climate change, will need to be agreed upon by all
parties and enforced collaboratively.
Finally, it is imperative that local people are actively involved
and/or employed in ongoing research, conservation, and education
projects, as it allows communities to attach a necessary personal
value to the conservation of their wildlife, but Oates (1999) and
Bergl et al. (2007) argued that community-based conservation projects increase pressure on protected areas and detract from overall
conservation goals. Rather, Oates (1999) suggested that government-sponsored conservation of nature for its intrinsic value alone,
supported by strict regulations and enforcement, can be successful. Given the current state of the BFH, the management of natural resources must seek to bridge the gap between conservation,
economic development, and human interests to ensure that the environment and the services it provides are not lost or overexploited
to the point of ecological collapse.
Acknowledgments
We would like to thank James Juvik, Stephanie Nagata, Donna Delparte, Jonathan Price, Sonia Juvik, Christoph Kueffer, and
all others involved in the organization of the Vulnerable Islands in
the Sky symposium. We would also like to thank Jose Manuel Esara Echube, Maximilliano Fero Meñe, Shaya Honarvar, Steve Woloszynek, Patrick McLaughlin, Jacob Owens, Demetrio Bocuma
Meñe, Erica Henn, Halle Choi, Manali Desai, Tessa Erickson, Joan
Taddei, and Laura Cronin for their valuable time and contributions.
References Cited
Abernethy, K. A., Coad, L., Taylor, G., Lee, M. E., and Maisels, F., 2013:
Extent and ecological consequences of hunting in Central African
rainforests in the twenty-first century. Philosophical Transactions
of the Royal Society B: Biological Sciences, 368: 20120303. http://
dx.doi.org/20120310.20121098/rstb.20122012.20120303.
Achard, F., Eva, H., Glinni, A., Mayaux, P., Richards, T., and Stibig, H.
J., 1998: Identification of Deforestation Hot Spot Areas in the Humid
Tropics. Ispra, Italy: Joint Research Centre, European Commission.
Albrechtsen, L., Fa, J. E., Barry, B., and Macdonald, D. W., 2006:
Contrasts in availability and consumption of animal protein in
Bioko Island, West Africa: the role of bushmeat. Environmental
Conservation, 32: 340–348.
Albrechtsen, L., Macdonald, D., Johnson, P. J., Castelo, R., and Fa, J.
E., 2007: Faunal loss from bushmeat hunting: empirical evidence
and policy implications in Bioko island. Environmental Science &
Policy, 10: 654–667.
Allen, C. D., Macalady, A. K., Chenchouni, H., Bachelet, D.,
McDowell, N., Vennetier, M., Kitzberger, T., Rigling, A., Breshears,
D. D., Hogg, E. H., Gonzalez, P., Fensham, R., Zhang, Z., Castro, J.,
Demidova, N., Lim, J.-H., Allard, G., Running, S. W., Semerci, A.,
and Cobb, N., 2010: A global overview of drought and heat-induced
tree mortality reveals emerging climate change risks for forests.
Forest Ecology and Management 259: 660–684.
Alpert, P., 1993: Conserving biodiversity in Cameroon. Ambio, 22:
44–49.
Amori, G., Gippoliti, S., and Helgen, K. M., 2008: Diversity,
distribution, and conservation of endemic island rodents. Quaternary
International, 182: 6–15. [cite this reference]
Anthony, N. M., Johnson-Bawe, M., Jeffery, K., Clifford, S. L.,
Abernethy, K. A., Tutin, C. E., Lahm, S. A., White, L. J. T., Utley,
J. F., Wickings, E. J., and Bruford, M. W., 2007: The role of
Pleistocene refugia and rivers in shaping gorilla genetic diversity in
central Africa. Proceedings of the National Academy of Sciences,
104: 20432–20436.
Assefa, A., Ehrich, D., Taberlet, P., Nemomissa, S., and Brochmann,
C., 2007: Pleistocene colonization of Afro-alpine ‘sky islands’ by
the Artic-alpine Arabis alpina. Heredity, 99: 133–142.
Barthlott, W., Lauer, W., and Placke, A., 1996: Global distribution
of species diversity in vascular plants: towards a world map of
phytodiversity. Erdkunde, 50: 317–328.
Beniston, M., 2000: Environmental change in mountains and uplands.
In Matthews, J. A., Bradley, R. S., Roberts, N., and Williams, M.
A. J. (eds.), Key Issues in Environmental Change. London: Hodder
Arnold.
Bennett, E. L., 2011: Another inconvenient truth: the failure of
enforcement systems to save charismatic species. Oryx, 45: 476–479.
Bergl, R. A., Oates, J. F., and Fotso, R., 2007: Distribution and
protected area coverage of endemic taxa in West Africa’s Biafran
forests and highlands. Biological Conservation, 134: 195–208.
Bergl, R. A., Bradley, B. J., Nsubuga, A., and Vigilant, L., 2008:
Effects of habitat fragmentation, population size and demographic
history on genetic diversity: the cross river gorilla in a comparative
context. American Journal of Primatology, 70: 848–859.
Bergl, R. A., Warren, Y., Nicholas, A., Dunn, A., Imong, I., SunderlandGroves, J., and Oates, J. F., 2011: Remote sensing analysis reveals
habitat, dispersal corridors and expanded distribution for the
critically endangered Cross River gorilla Gorilla gorilla diehli.
Oryx, 46: 278–289.
Blackburn, D. C., 2010: A new squeaker frog (Arthroleptidae:
Arthroleptis) from Bioko Island, Equatorial Guinea. Herpetologica,
66: 320–334.
Blom, A., Yamindou, J., and Prins, H. H. T., 2004: Status of the protected
areas of the Central African Republic. Biological Conservation, 118:
479–487.
Bonnefille, R., Roeland, J. C., and Guiot, J., 1990: Temperature and
rainfall estimates for the past 40,000 years in equatorial Africa.
Nature, 346: 347–349.
Brashares, J. S., Arcese, P., and Sam, M. K., 2001: Human demography
and reserve size predict wildlife extinction in West Africa.
Proceedings of the Royal Society of London Series B–Biological
Sciences, 268: 2473–2478.
Brooks, T., Balmford, A., Burgess, N., Fjeldså, J. O. N., Hansen, L. A.,
Moore, J., Rahbek, C., and Williams, P., 2001: Toward a blueprint
for conservation in Africa. Bioscience, 51: 613–624.
Bruner, A. G., Gullison, R. E., Rice, R. E., and da Fonseca, G. A. B.,
2001: Effectiveness of parks in protecting tropical biodiversity.
Science, 291: 125–128.
Buchanan, G. M., Donald, P. F., and Butchart, S. H. M., 2011:
Identifying priority areas for conservation: a global assessment
for forest-dependent birds. PLoS One, 6: e29080, http://dx.doi.
org/10.1371/journal.pone.0029080.
Burke, K., 2001: Origin of the Cameroon Line of volcano-capped
swells. Journal of Geology, 109: 349–362.
Bush, M. B., 2002: Distributional change and conservation on the
Andean flank: a palaeoecological perspective. Global Ecology and
Biogeography, 11: 463–473.
Butynski, T. M., and Koster, S. H., 1994. Distribution and conservation
status of primates in Bioko Island, Equatorial Guinea. Biodiversity
and Conservation, 3: 893–909.
Butynski, T. B., Schaaf, C. D., and Hearn, G. W., 1997: African Buffalo
Syncerus caffer extirpated on Bioko Island, Equatorial Guinea.
Journal of African Zoology, 111: 57–61.
Cable, S., and Cheek, M., 1998: The Plants of Mt. Cameroon: a
Conservation Checklist. Royal Botanical Gardens, Kew.
Campbell, G., Kuehl, H., Diarrassouba, A., N’Goran, P. K., and
Boesch, C., 2011: Long-term research sites as refugia for threatened
and over-harvested species. Biology Letters, 7: 723–726.
Cerutti, P. O., Nasi, R., and Tacconi, L., 2008: Sustainable forest
management in Cameroon needs more than approved forest
management plans. Ecology and Society, 13: 36.
Chapman, C. A., and Onderdonk, D. A., 1998: Forests without primates:
Primate/plant codependency. American Journal of Primatology, 45:
127–141.
Chapman, H. M., Olson, S. M., and Trum, D., 2004: An assessment of
changes in the montane forests of Taraba State, Nigeria, over the past
30 years. Oryx, 38: 282–290.
Charlotte, C. N., 2010: Cadre Fonctionnel de Gestion du Parc National du
Mont Cameroun. Vol. 2 of Cameroon—Competitive Value Chains Project:
Environment and Social Management Plan. Ministere de L’Économie, de
la Planification et de L’Amenagement du Territoire, Cameroon.
Chen, I.-C., Shiu, H.-J., Benedick, S., Holloway, J. D., Chey, V. K.,
Barlow, H. S., Hill, J. K., and Thomas, C. D., 2009: Elevation
increases in moth assemblages over 42 years on a tropical mountain.
Proceedings of the National Academy of Sciences, 106: 1479–1483.
Chen, I.-C., Hill, J. K., Ohlemüller, R., Roy, D. B., and Thomas, C. D.,
2011: Rapid range shifts of species associated with high levels of
climate warming. Science, 333: 1024–1026.
CIA, 2013: The World Fact Book 2013. Washington DC: Central
Intelligence Agency. Available from https://www.cia.gov/library/
publications/the-world-factbook/index.html.
Colwell, R. K., Brehm, G., Cardelús, C. L., Gilman, A. C., and
Longino, J. T., 2008: Global warming, elevational range shifts,
and lowland biotic attrition in the wet tropics. Science, 322:
258–261.
Cronin, D. T. 2013. The Impact of Bushmeat Hunting on the Primates
of Bioko Island, Equatorial Guinea. Ph.D. thesis, Department of
Biology, Drexel University, Philadelphia, Pennsylvania.
Cronin, D. T., Bocuma Meñe, D., Butynski, T. B., Echube, J. M. E.,
Hearn, G. W., Honarvar, S., Owens, J. R., and Bohome, C. P., 2010:
Opportunities Lost: The Rapidly Deteriorating Conservation Status
of the Monkeys on Bioko Island, Equatorial Guinea. A report to
the government of Equatorial Guinea by the Bioko Biodiversity
Protection Program, Drexel University, Philadelphia, Pennsylvania.
Cronin, D. T., Riaco, C., and Hearn, G. W., 2013: Survey of threatened
monkeys in the Iladyi River Valley Region, Southeastern Bioko
Island, Equatorial Guinea. African Primates, 8: 1–8.
DREW T. CRONIN ET AL. / 901
deMenocal, P. B., 1995: Plio-Pleistocene African climate. Science,
270: 53–59.
de Merode, E., and Cowlishaw, G., 2006: Species protection, the
changing informal economy, and the politics of access to the
bushmeat trade in the Democratic Republic of Congo. Conservation
Biology, 20: 1262–1271.
de Wasseige, C., de Marcken, P., Bayol, N., Hiol Hiol, F., Mayaux, P.,
Desclée, B., Nasi, R., Billand, A., Defourny, P., and Eba’a Atyi, R.
(eds.), 2012: The Forests of the Congo Basin—State of the Forest
2010. Luxembourg: Publications Office of the European Union.
Deruelle, B., Moreau, C., Nkoumbou, C., Kambou, R., Lissom, J.,
Njongfang, E., Ghogomu, R. T., and Nono, A., 1991: The Cameroon
Line: a review. In Kampunzu, A. B., and Lubala, R. T. (eds.),
Magmatism in Extensional Structural Settings. Berlin: Springer
Verlag, 274–327.
Djeukam, R., Ntolo, M., Dinga, N., Tedjiozem, R., Talla, M., and Njike,
H., 2012: The Wildlife Law as a Tool for Protecting Threatened
Species in Cameroon. Cameroon: Ministry of Forestry and Wildlife
(MINFOF), Department of Wildlife and Protected Areas.
Dudley, N. (ed.), 2008: Guidelines for Applying Protected Area
Management Categories. Gland, Switzerland: IUCN.
Eggermont, H., Verschuren, D., Audenaert, L., Lens, L., Russell, J.,
Klaassen, G., and Heiri, O., 2010: Limnological and ecological
sensitivity of Rwenzori mountain lakes to climate warming.
Hydrobiologia, 648: 123–142.
Eisentraut, M., 1973: Die Wirbeltierfauna von Fernando Poo und West
Kamerun. Bonner Zoologische Monographien, 3: 1–428.
Exell, A. W., 1973: Angiosperms of the islands of the Gulf of Guinea
(Fernando Po, Príncipe, São Tomé and Annobón). Bulletin of the
British Museum (Natural History), Botany, 4: 325–411.
Fa, J. E., 2000: Hunted animals in Bioko Island, West Africa:
sustainability and future. In Robinson, J. G., and Bennett, E. L.
(eds.), Hunting for Sustainability in Tropical Forests. New York:
Columbia University Press.
Fa, J. E., Yuste, J. E. G., and Castelo, R., 2000: Bushmeat markets
on Bioko Island as a measure of hunting pressure. Conservation
Biology, 14: 1602–1613.
Fa, J. E., Seymour, S., Dupain, J. E. F., Amin, R., Albrechtsen, L.,
and Macdonald, D., 2006: Getting to grips with the magnitude of
exploitation: bushmeat in the Cross-Sanaga Rivers region, Nigeria
and Cameroon. Biological Conservation, 129: 497–510.
Figueiredo, E., 1994: Diversity and endemism of angiosperms in the
Gulf of Guinea islands. Biodiversity and Conservation, 3: 785–793.
Fischer, F., 2008: The importance of law enforcement for protected
areas: Don’t Step Back! Be Honest—Protect! GAIA—Ecological
Perspectives for Science and Society, 17: 101–103.
Flenley, J. R., 1979: The Equatorial Rain Forest: A Geological History.
Boston: Butterworths Publishers.
Forboseh, P. F., Sunderland, T. C. H., Comiskey, J. A., and Balinga,
M., 2011: Tree population dynamics of three altitudinal vegetation
communities on Mount Cameroon (1989–2004). Journal of
Mountain Science, 8: 495–504.
Fotso, R., Dowsett-Lemaire, F., Dowsett, R. J., Cameroon
Ornithological Club, Scholte, P., Languy, M., and Bowden, C., 2001:
Important bird areas in Africa and associated islands—Cameroon.
In Fishpool, D. C., and Evans, M. I. (eds.), Important Bird Areas
in Africa and Associated Islands: Priority Sites for Conservation.
Newbury and Cambridge, UK: Birdlife International, 133–159.
Gartshore, M. E., 1984: The status of montane herpetofauna of
the Cameroon Highlands. In Stuart, S. N. (ed.), Conservation of
Cameroon Montane Forests. Cambridge: International Council for
Bird Preservation, 204–240.
Gottelli, D., Marino, J., Sillero-Zubiri, C., and Funk, S. M., 2004: The
effect of the last glacial age on speciation and population genetic
structure of the endangered Ethiopian wolf (Canis simensis).
Molecular Ecology, 13: 2275–2286.
Green, A. A., and Rodewald, P. G., 1996: New bird records from Korup
National Park and environs, Cameroon. Malimbus, 18: 122–133.
902 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH
Haffer, J., 1969: Speciation in Amazonian forest birds. Science, 165:
131–137.
Hansen, M. C., Stehman, S. V., Potapov, P. V., Loveland, T. R.,
Townshend, J. R. G., DeFries, R. S., Pittman, K. W., Arunarwati,
B., Stolle, F., Steininger, M. K., Carroll, M., and DiMiceli, C., 2008:
Humid tropical forest clearing from 2000 to 2005 quantified by using
multitemporal and multiresolution remotely sensed data. Proceedings
of the National Academy of Sciences, 105(27): 9439–9444.
Hart, T. B., Hart, J. A., and Murphy, P. G., 1989: Monodominant
and species-rich forests of the humid tropics: causes for their cooccurrence. American Naturalist, 133: 613–633.
Hilborn, R., Arcese, P., Borner, M., Hando, J., Hopcraft, G., Loibooki,
M., Mduma, S., and Sinclair, A. R. E., 2006: Effective enforcement
in a conservation area. Science, 314: 1266.
Holmern, T., Muya, J., and Røskaft, E., 2007: Local law enforcement
and illegal bushmeat hunting outside the Serengeti National Park,
Tanzania. Environmental Conservation, 34: 55–63.
Huarez, B., Petre, C.-A., and Doucet, J.-L., 2013: Impacts of logging
and hunting on western lowland gorilla (Gorilla gorilla gorilla)
populations and consequences for forest regeneration. A review.
Biotechnology, Agronomy, Society and Environment, 7: 364–372.
Ingram, V., and Nsawir, A. T., 2007: Pygeum: money growing on trees
in the Cameroon highlands? Nature & Faune, 22: 29–36.
IPCC, 2014a: Part A: global and sectoral aspects. In Field, C. B.,
Barros, V. R., Dokken, D. J., Mach, K. J., Mastrandrea, M. D., Bilir,
T. E., Chatterjee, M., Ebi, K. L., Estrada, Y. O., Genova, R. C.,
Girma, B., Kissel, E. S., Levy, A. N., MacCracken, S., Mastrandrea,
P. R., and White, L. L. (eds.), Climate Change 2014: Impacts,
Adaptation, and Vulnerability. Contribution of Working Group II
to the Fifth Assessment Report of the Intergovernmental Panel on
Climate Change. Cambridge: Cambridge University Press.
IPCC, 2014b: Part B: regional aspects. In Barros, V. R., Field, C.
B., Dokken, D. J., Mastrandrea, M. D., Mach, K. J., Bilir, T. E.,
Chatterjee, M., Ebi, K. L., Estrada, Y. O., Genova, R. C., Girma,
B., Kissel, E. S., Levy, A. N., MacCracken, S., Mastrandrea, P. R.,
and White, L. L. (eds.), Climate Change 2014: Impacts, Adaptation,
and Vulnerability. Contribution of Working Group II to the Fifth
Assessment Report of the Intergovernmental Panel on Climate
Change. Cambridge: Cambridge University Press.
IUCN, 2013: IUCN Red List of Threatened Species. Version 2013.1.
www.iucnredlist.org, accessed 1 August 2013.
IUCN and UNEP, 2010: The World Database on Protected Areas
(WDPA). Cambridge, UK: UNEP-WCMC.
Jachmann, H., 2008: Monitoring law-enforcement performance in nine
protected areas in Ghana. Biological Conservation, 141: 89–99.
James, A. N., Green, M. J. B., and Paine, J. R., 1999: Global Review
of Protected Area Budgets and Staff. Cambridge, UK: World
Conservation Monitoring Center.
James, R., Washington, R., and Rowell, D. P., 2013. Implications of
global warming for the climate of African rainforests. Philosophical
Transactions of the Royal Society B: Biological Sciences, 368: http://
dx.doi.org/10.1098/rstb.2012.0298.
Jones, P. J., 1994: Biodiversity in the Gulf of Guinea—an overview.
Biodiversity and Conservation, 3: 772–784.
Kingdon, J., 1997: The Kingdon Field Guide to African Mammals. San
Diego: Academic Press.
Laporte, N. T., Stabach, J. A., Grosch, R., Lin, T. S., and Goetz, S.
J., 2007: Expansion of industrial logging in Central Africa. Science,
316: 1451.
Laurance, W. F., Goosem, M., and Laurance, S. G. W., 2009: Impacts
of roads and linear clearings on tropical forests. Trends in Ecology
& Evolution, 24: 659–669.
Laurance, W. F., Carolina Useche, D., Shoo, L. P., Herzog, S. K., Kessler,
M., Escobar, F., Brehm, G., Axmacher, J. C., Chen, I. C., Gámez, L. A.,
Hietz, P., Fiedler, K., Pyrcz, T., Wolf, J., Merkord, C. L., Cardelus, C.,
Marshall, A. R., Ah-Peng, C., Aplet, G. H., del Coro Arizmendi, M.,
Baker, W. J., Barone, J., Brühl, C. A., Bussmann, R. W., Cicuzza, D.,
Eilu, G., Favila, M. E., Hemp, A., Hemp, C., Homeier, J., Hurtado, J.,
Jankowski, J., Kattán, G., Kluge, J., Krömer, T., Lees, D. C., Lehnert,
M., Longino, J. T., Lovett, J., Martin, P. H., Patterson, B. D., Pearson,
R. G., Peh, K. S. H., Richardson, B., Richardson, M., Samways, M. J.,
Senbeta, F., Smith, T. B., Utteridge, T. M. A., Watkins, J. E., Wilson,
R., Williams, S. E., and Thomas, C. D., 2011: Global warming,
elevational ranges and the vulnerability of tropical biota. Biological
Conservation, 144: 548–557.
Lawson, D. P., 1993: The reptiles and amphibians of the Korup National
Park Project, Cameroon. Herpetological Natural History, 1: 27–90.
Leuschner, C., 1996: Timberline and alpine vegetation on the tropical
and warm-temperate oceanic islands of the world: elevation,
structure and floristics. Vegetatio, 123: 193–206.
Linder, J. M., 2013: African primate diversity threatened by “new
wave” of industrial oil palm expansion. African Primates, 8: 25–38.
Linder, J. M., and Oates, J. F., 2011: Differential impact of bushmeat
hunting on monkey species and implications for primate conservation
in Korup National Park, Cameroon. Biological Conservation, 144:
738–745.
Maisels, F., Keming, E., Kemei, M., and Toh, C., 2001: The extirpation
of large mammals and implications for montane forest conservation:
the case of the Kilum-Ijim Forest, North-west Province, Cameroon.
Oryx, 35: 322–331.
Maley, J., Livingstone, D. A., Giresse, P., Thouveny, N., Brenac, P.,
Kelts, K., Kling, G., Stager, C., Haag, M., Fournier, M., Bandet,
Y., Williamson, D., and Zogning, A., 1990: Lithostratigraphy,
volcanism, paleomagnetism and palynology of Quaternary lacustrine
deposits from Barombi Mbo (West Cameroon): preliminary results.
Journal of Volcanology and Geothermal Research, 42: 319–335.
Marzoli, A., Piccirillo, E. M., Renne, P. R., Bellieni, G., Iacumin, M.,
Nyobe, J. B., and Tongwa, A. T., 2000: The Cameroon Volcanic
Line revisited: petrogenesis of continental basaltic magmas
from lithospheric and asthenospheric mantle sources. Journal of
Petrology, 41: 87–109.
Megevand, C., Mosnier, A., Hourticq, J., Sanders, K., Doetinchem,
N., and Streck, C., 2013: Deforestation Trends in the Congo Basin:
Reconciling Economic Growth and Forest Protection. Washington,
D.C.: The World Bank.
Moreau, R. E., 1963: Vicissitudes of the African biomes in the late
Pleistocene. Proceedings of the Zoological Society of London, 141:
395–421.
Morra, W., Hearn, G., and Buck, A. J., 2009: The market for bushmeat:
Colobus satanas on Bioko Island. Ecological Economics, 68: 2619–
2626.
Myers, N., Mittermeier, R. A., Mittermeier, C. G., da Fonseca, G.
A. B., and Kent, J., 2000: Biodiversity hotspots for conservation
priorities. Nature, 403: 853–858.
Newmark, W. D., 2008: Isolation of African protected areas. Frontiers
in Ecology and the Environment, 6: 321–328.
N’Goran, P. K., Boesch, C., Mundry, R., N’Goran, E. K., Herbinger,
I., Yapi, F. A., and Kühl, H. S., 2012: Hunting, law enforcement, and
African primate conservation. Conservation Biology, 26: 565–571.
Nguiffo, S., and Talla, M., 2010: Cameroon’s wildlife legislation: local
custom versus legal conception. Unasylva, 61: 14–18.
Njuh Fuo, O., and Memuna Semi, S., 2011: Cameroon’s environmental
framework law and the balancing of interests in socio-economic
development. In Faure, M., and de Plessis, W. (eds.), The Balancing
of Interests in Environmental Law in Africa. Pretoria, South Africa:
Pretoria University Law Press.
Nosti, J., 1947: Notas geograficas, fisicas y economicas sobre los
territorios espanoles del Golfo de Guinea. Consejo Superior de
Investigaciones Cientificas, Instituto de Estudios Africanos, Madrid.
Oates, J. F., 1996: African Primates: Status Survey and Conservation
Action Plan. Gland, Switzerland: International Union for
Conservation of Nature (IUCN) and Species Survival Commission
(SSC), Primate Specialist Group.
Oates, J. F., 1999: Myth and Reality in the Rain Forest: How
Conservation Strategies Are Failing in West Africa. Berkeley:
University of California Press.
Oates, J. F., 2011: Primates of West Africa: A Field Guide and Natural
History. Arlington, Virginia: Conservation International.
Oates, J. F., Bergl, R. A., and Linder, J. M., 2004: Africa’s Gulf of
Guinea Forests: Biodiversity Patterns and Conservation Priorities.
Advances in Applied Biodiversity Science, Volume 6. New York:
Wildlife Conservation Society (WCS), and Washington, D.C.: Center
for Applied Biodiversity Science (CABS), Conservation International.
Ohlemüller, R., Anderson, B. J., Araújo, M. B., Butchart, S. H.
M., Kudrna, O., Ridgely, R. S., and Thomas, C. D., 2008: The
coincidence of climatic and species rarity: high risk to small-range
species from climate change. Biology Letters, 4: 568–572.
Olson, D. M., Dinerstein, E., Wikramanayake, E. D., Burgess, N.
D., Powell, G. V. N., Underwood, E. C., D’Amico, J. A., Itoua, I.,
Strand, H. E., Morrison, J. C., Loucks, C. J., Allnutt, T. F., Ricketts,
T. H., Kura, Y., Lamoreux, J. F., Wettengel, W. W., Hedao, P., and
Kassem, K. R., 2001: Terrestrial ecoregions of the world: a new map
of life on Earth. Bioscience, 51: 933–938.
Onana, J. M., and Cheek, M., 2011: Red Data Book of the Flowering
Plants of Cameroon: IUCN Global Assessments. Kew, U.K.: Royal
Botanic Gardens.
Payton, R. W., 1993: Ecology, Altitudinal Zonation and Conservation
of Tropical Rainforest of Mt. Cameroon. Report to ODA, London.
Peh, K. S. H., and Drori, O., 2010: Fighting corruption to save the
environment: Cameroon’s experience. AMBIO: A Journal of the
Human Environment, 39: 336–339.
Penlap, E. K., Matulla, C., von Storch, H., and Kamga, F. M., 2004:
Downscaling of GCM scenarios to assess precipitation changes
in the little rainy season (March–June) in Cameroon. Climate
Research, 26: 85–96.
Pérez del Val, J., Fa, J. E., Castroviejo, J., and Purroy, F. J., 1994:
Species richness and endemism of birds in Bioko. Biodiversity and
Conservation, 3: 868–892.
Pimentel, D., McNair, M., Buck, L., Pimentel, M., and Kamil, J., 1997:
The value of forests to world food security. Human Ecology, 25: 91–120.
Pounds, J. A., Fogden, M. P. L., and Campbell, J. H., 1999: Biological
response to climate change on a tropical mountain. Nature, 398:
611–615.
Pounds, J. A., Bustamante, M. R., Coloma, L. a., Consuegra, J.
a., Fogden, M. P. L., Foster, P. N., La Marca, E., Masters, K. L.,
Merino-Viteri, A., Puschendorf, R., Ron, S. R., Sánchez-Azofeifa,
G. A., Still, C. J., and Young, B. E., 2006: Widespread amphibian
extinctions from epidemic disease driven by global warming.
Nature, 439: 161–167.
Qorvis, 2010: Equatorial Guinea President Pledges Environmental
Conservation. Qorvis Communications, www.prnewswire.
com/news-releases/equatorial-guinceruea-president-pledgesenvironmental-conservation-97605059.html.
Raxworthy, C. J., Pearson, R. G., Rabibisoa, N., Rakotondrazafy, A.
M., Ramanamanjato, J.-B., Raselimanana, A. P., Wu, S., Nussbaum,
R. A., and Stone, D. A., 2008: Extinction vulnerability of tropical
montane endemism from warming and upslope displacement: a
preliminary appraisal for the highest massif in Madagascar. Global
Change Biology, 14: 1703–1720.
Redford, K. H., 1992: The empty forest. Bioscience, 42: 412–422.
Republic of Cameroon, 1994: Law No. 94-01 of 20 January 1994: To
Lay Down Forestry, Wildlife, and Fisheries Regulations, Republic of
Cameroon, Yaounde, Cameroon.
Republic of Cameroon, 1995: Decree No. 95-466-PM of 20 July 1995:
To Lay Down the Conditions for the Implementation of Wildlife
Regulations, Republic of Cameroon, Yaounde, Cameroon.
Republic of Cameroon, 1996: Law No. 96-12 of 5 August 1996:
Relating to Environmental Management, Republic of Cameroon,
Yaounde, Cameroon.
Republic of Equatorial Guinea, 1988: Regulation of Wildlife, Hunting,
and Protected Areas. Law number 8/1998, Malabo, Republic of
Equatorial Guinea.
Republic of Equatorial Guinea, 2000: Protected Areas Law. Law
number 4/2000, Malabo, Republic of Equatorial Guinea.
DREW T. CRONIN ET AL. / 903
Republic of Equatorial Guinea, 2003: Environmental Regulation Law
in the Republic of Equatorial Guinea. Law number 7/2003, Republic
of Equatorial Guinea.
Republic of Equatorial Guinea, 2007: Hunting and Consumption of
Monkeys and Other Primates in the Republic of Equatorial Guinea
Is Prohibited. Law number 72/2007, Republic of Equatorial Guinea.
Richards, P. W., 1963: Ecological notes on West African vegetation
III. The upland forests of Cameroons Mountain. Journal of Ecology,
51: 529–554.
Richards, P. W., 1996: The Tropical Rain Forest: An Ecological Study.
Cambridge: Cambridge University Press.
Robinson, J. G., and Bennett, E. L. (eds.), 2000: Hunting for
Sustainability in Tropical Forests. New York: Columbia University
Press.
Rowcliffe, J. M., de Merode, E., and Cowlishaw, G., 2004: Do wildlife
laws work? Species protection and the application of a prey choice
model to poaching decisions. Proceedings of the Royal Society of
London. Series B: Biological Sciences, 271: 2631–2636.
Roy, M. S., 1997: Recent diversification in African greenbuls
(Pycnonotidae: Andropadus) supports a montane speciation model.
Proceedings of the Royal Society of London. Series B: Biological
Sciences, 264: 1337–1344.
Sanderson, E. W., Jaiteh, M., Levy, M. A., Redford, K. H., Wannebo,
A. V., and Woolmer, G., 2002: The human footprint and the last of
the wild. Bioscience, 52: 891–904.
Schiotz, A., 1999: Treefrogs of Africa. Frankfurt: Edition Chimaira.
Schloss, C. A., Nuñez, T. A., and Lawler, J. J., 2012: Dispersal will
limit ability of mammals to track climate change in the Western
Hemisphere. Proceedings of the National Academy of Sciences, 109:
8606–8611.
Sekercioglu, C. H., Schneider, S. H., Fay, J. P., and Loarie, S. R.,
2008: Climate change, elevational range shifts, and bird extinctions.
Conservation Biology, 22: 140–150.
Smith, T. B., Holder, K., Girman, D., O’Keefe, K., Larison, B., and
Chan, Y., 2000: Comparative avian phylogeography of Cameroon
and Equatorial Guinea mountains: implications for conservation.
Molecular Ecology, 9: 1505–1516.
Smith, R. J., Muir, R. D., Walpole, M. J., Balmford, A., and LeaderWilliams, N., 2003: Governance and the loss of biodiversity. Nature,
426: 67–70.
Stattersfield, A. J., Crosby, M. J., Long, A. J., and Wege, D. C.,
1998: Endemic Bird Areas of the World: Priorities for Biodiversity
Conservation. Cambridge: Birdlife International.
Still, C. J., Foster, P., and Schneider, S., 1999: Simulating the effects
of climate change on tropical montane cloud forests. Nature, 398:
608–610.
Struhsaker, T. T., Struhsaker, P. J., and Siex, K. S., 2005: Conserving
Africa’s rain forests: problems in protected areas and possible
solutions. Biological Conservation, 123: 45–54.
Suh, C. E., Sparks, R. S. J., Fitton, J. G., Ayonghe, S. N., Annen, C.,
Nana, R., and Luckman, A., 2003: The 1999 and 2000 eruptions of
Mount Cameroon: eruption behaviour and petrochemistry of lava.
Bulletin of Volcanology, 65: 267–281.
SWPDFW, GTZ–Programme for the Sustainable Management of
Natural Resources, and WWF Coastal Forests Programme, 2005:
Technical Note for the Creation of the Mount Cameroon National
Park. Buea, Cameroon: GTZ.
Tchouto, P., Edwards, I., Cheek, M., Ndam, N., and Acworth, J., 1999:
Mount Cameroon Cloud Forest. In Timberlake, J., and Kativu, S.
(eds.), African Plants: Biodiversity, Taxonomy, and Uses. Kew,
U.K.: Royal Botanic Gardens, 263–277.
Thomas, C. D., Cameron, A., Green, R. E., Bakkenes, M., Beaumont,
L. J., Collingham, Y. C., Erasmus, B. F. N., de Siqueira, M. F.,
Grainger, A., Hannah, L., Hughes, L., Huntley, B., van Jaarsveld,
904 / ARCTIC, ANTARCTIC, AND ALPINE RESEARCH
A. S., Midgley, G. F., Miles, L., Ortega-Huerta, M. A., Townsend
Peterson, A., Phillips, O. L., and Williams, S. E., 2004: Extinction
risk from climate change. Nature, 427: 145–148.
Tranquilli, S., Abedi-Lartey, M., Amsini, F., Arranz, L., Asamoah,
A., Babafemi, O., Barakabuye, N., Campbell, G., Chancellor, R.,
Davenport, T. R. B., Dunn, A., Dupain, J., Ellis, C., Etoga, G.,
Furuichi, T., Gatti, S., Ghiurghi, A., Greengrass, E., Hashimoto, C.,
Hart, J., Herbinger, I., Hicks, T. C., Holbech, L. H., Huijbregts, B.,
Imong, I., Kumpel, N., Maisels, F., Marshall, P., Nixon, S., Normand,
E., Nziguyimpa, L., Nzooh-Dogmo, Z., Okon, D. T., Plumptre, A.,
Rundus, A., Sunderland-Groves, J., Todd, A., Warren, Y., Mundry,
R., Boesch, C., and Kuehl, H., 2012: Lack of conservation effort
rapidly increases African great ape extinction risk. Conservation
Letters, 5: 48–55.
Tsafack, J. F., Wandji, P., Bardintzeff, J., Bellon, H., and Guillou, H.,
2009: The Mount Cameroon stratovolcano (Cameroon Volcanic
Line, Central Africa): Petrology, geochemistry, isotope, and age
data. Geochemistry, Mineralogy and Petrology, 47: 65–78.
Tye, H., 1984: Geology and landforms in the highlands of western
Cameroon. In Stuart, S. N. (ed.), Conservation of Cameroon
Montane Forests. Cambridge, U.K.: International Council for Bird
Preservation, 15–17.
UNDP-GEF, 2010: Strengthening the national system of protected
areas in Equatorial Guinea for the effective conservation of
representative ecosystems and globally significant biodiversity.
United Nations Development Programme, Global Environmental
Fund, Project Report No. 4185.
Vanthomme, H., Belle, B., and Forget, P. M., 2010: Bushmeat hunting
alters recruitment of large-seeded plant species in Central Africa.
Biotropica, 42: 672–679.
Wang, B. C., Sork, V. L., Leong, M. T., and Smith, T. B., 2007: Hunting
of mammals reduces seed removal and dispersal of the Afrotropical
tree Antrocaryon klaineanum (Anacardiaceae). Biotropica, 39: 340–
347.
Wilkie, D. S., Bennett, E. L., Peres, C. A., and Cunningham, A. A.,
2011: The empty forest revisited. Annals of the New York Academy
of Sciences, 1223: 120–128.
Willis, K. J., Bennett, K. D., Burrough, S. L., Macias-Fauria, M.,
and Tovar, C., 2013: Determining the response of African biota to
climate change: using the past to model the future. Philosophical
Transactions of the Royal Society B: Biological Sciences, 368: http://
dx.doi.org/10.1098/rstb.2012.0491.
Wittemyer, G., Elsen, P., Bean, W. T., Coleman, A., Burton, O., and
Brashares, J. S., 2008: Accelerated human population growth at
protected area edges. Science, 321: 123–126.
Wolfe, N. D., Daszak, P., Marm Kilpatrick, A., and Burke, D. S., 2005:
Bushmeat hunting, deforestation, and prediction of zoonotic disease
emergence. Emerging Infectious Diseases, 11: 1822–1827.
World Bank, 2014a: Cameroon—GDP (current US Dollars). The World
Bank, http://data.worldbank.org/country/cameroon?display=default.
World Bank, 2014b: Equatorial Guinea—GDP (current US Dollars).
The World Bank, http://data.worldbank.org/country/equatorialguinea?display=default.
Zimkus, B. M., 2009: Biogeographical analysis of Cameroonian
puddle frogs and description of a new species of Phrynobatrachus
(Anura: Phrynobatrachidae) endemic to Mount Oku, Cameroon.
Zoological Journal of the Linnean Society, 157: 795–813.
Zofou, D., Kowa, T. K., Wabo, H. K., Ngemenya, M. N., Tane, P., and
Titanji, V. P. K., 2011: Hypericum lanceolatum (Hypericaceae) as
a potential source of new anti-malarial agents: a bioassay-guided
fractionation of the stem bark. Malaria Journal, 10: 1–7.
MS accepted August 2014