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Short Communication
African development corridors intersect key
protected areas
Sean Sloan
1
*, Bastian Bertzky
2,3
and
William F. Laurance
1,2
1
Center for Tropical Environmental and Sustainability Science,
College of Marine and Environmental Sciences, James Cook
University, Cairns, Qld 4870, Australia,
2
European
Commission, Joint Research Centre (JRC), Institute for
Environment and Sustainability (IES), Via Enrico Fermi
2749, 21027 Ispra, VA, Italy and
3
International Union for
Conservation of Nature (IUCN), World Heritage Programme,
Rue Mauverney 28, 1196 Gland, Switzerland
Introduction
Unprecedented growth of mining and agriculture in Africa
is encroaching on remaining habitats. Mining in Africa
frequently occurs in proximity to protected areas (PAs),
more than in other world regions (Dur
an, Rauch & Gaston,
2013), and at least 23 African PAs have been degazetted
or downgraded as a result (Edwards et al., 2014). Even
natural World Heritage Sites, the global pinnacle of
conservation, suffer mining and oil/gas exploration and
exploitation across 31 sites and 18 African countries
(WWF, 2015), again more than other world regions
(WWF, 2016). The anticipated expansion of transport
‘development corridors’ related to infrastructure and
resource development could impact the ecological integrity
of many other PAs as roads and rails link producers with
refineries and ports over vast distances (Weng et al.,
2013).
Recent analysis highlights frequent intersections
between 33 African development corridors and PAs
(Laurance et al., 2015b). Only five existing or proposed
corridors were considered ‘advisable’ given their potential
environmental costs and agricultural benefits (another 22
were ‘marginal’ and six ‘inadvisable’). However, while
insightful, the aggregate analysis of Laurance et al.
(2015b) overlooked important differences between the
conservation objectives of intersected PAs. Protected-area
significance and management objectives vary along a
spectrum from global importance and/or strict protection
to local importance and nominally regulated exploitation.
Only half of the 15% of African PAs with reported IUCN
management categories had stricter protection levels
(Categories I-IV) indicative of greater levels of natural
integrity –a low figure by global standards (Deguignet
et al., 2014, IUCN and UNEP-WCMC, 2015). It remains
unclear which pole of this ‘PA spectrum’ is most threat-
ened by corridors and to which degree. This uncertainty is
compounded by the fact that most of the 33 corridors are
in some stage of planning, so that it is also unclear
whether the greatest impacts are pending or already
realized. The ongoing expansion of Africa’s development
corridors and mining sector urges the resolution of these
uncertainties to inform regional environmental planning
(Edwards et al., 2014; Laurance et al., 2014).
Here, we detail actual and potential intersections
between African development corridors and a range of
PA designations, focusing on designations of the greatest
conservation significance, to clarify corridors’ potential
effects on conservation objectives. We further detail
variations in corridors’ intersections with PAs according
to their development status in order to differentiate
between current and potential future ecological impacts.
The actual effect of intersections on ecological integrity is
inevitably site specific and, in special circumstances, might
entail benefits; yet, the weight of historical experience
suggests that intersections by transportation infrastructure
often pose challenges to ecological integrity insofar as they
facilitate deforestation, logging or resource extraction,
poaching, fragmentation of ecological communities, or
hydrological and climatic changes in PAs (Forman &
Alexander, 1998; Wilkie et al., 2000; Walsh et al., 2003;
Blake et al., 2007, 2008; Laurance, Goosem & Laurance,
2009; Freitas, Hawbaker & Metzger, 2010; Maisels et al.,
2013; Barber et al., 2014; Perz, 2014; Hopcraft et al.,
2015a). Our analysis shows that corridors, and particu-
larly proposed corridors, disproportionately intersect, and
thus by extension may challenge the ecological integrity of
the most important PAs in Africa.
*Correspondence: E-mail: sean.sloan@jcu.edu.au
©2016 John Wiley & Sons Ltd, Afr. J. Ecol. 1
Materials and methods
PA boundaries for this analysis come from the World
Database on Protected Areas (WDPA) (IUCN and UNEP-
WCMC, 2015). We focused on four PA designations
generally reserved for areas of special conservation value:
World Heritage Sites (WHS), Ramsar Sites, UNESCO Man-
and-Biosphere (MAB) Reserves and National Parks. These
designations are hereafter termed ‘key PAs’. The first three
are international conservation designations, and the latter
is the most important national designation for tourism
(Dudley, 2008). These designations are applied relatively
consistently across countries, facilitating continental-scale
analysis. We also considered national-level PA designa-
tions beyond National Parks but grouped these themati-
cally (Table 1; Table S1) because they are not standardized
amongst countries. It was not possible to similarly analyse
IUCN management categories because 85% of African PAs
lack this information in the WDPA. PAs here are defined as
unique WDPA records distinguishing distinct spatio-
administrative entities, each with distinct designations,
conservation objectives and management regimes.
On some occasions, the WDPA enumerates part or all of
a given area twice with each entry assigned a distinct PA
designation. For example, a communal forest reserve may
partially overlap a national park, or a portion of a national
park may also be a WHS. While such entities constitute
two PAs both in the WDPA and administratively in
practice, this classification scheme slightly exaggerates
estimates of the number of PAs intersected by corridors as
well as of the total length/area of PA–corridor intersections
relative to scenarios overlooking PA overlap
(Appendix S1). We controlled for this in two ways to
ensure robust and meaningful generalizations. First, we
compared PA-designation frequencies between intersected
PAs and all African PAs. Any exaggeration in the
frequencies of intersected PA designations should be
proportional to that of the set of all African PAs if
corridors intersect PA designations without bias. Thus,
any significant difference in PA-designation frequencies
between these two sets of PAs would signal corridors’ bias
in intersecting PAs of particular designations. Comparisons
considered the four key PA designations individually but
collapsed groups of national-level PA designations
(Table 1) into one group because their individual frequen-
cies could not be determined in the very large set of all
African PAs (n =6,546). Second, to complement the
estimates of the total length/area of PA–corridor
intersections for all PA designations, we also present
estimates of total length/area overlooking PA overlaps
(Table 1). The latter ‘aggregate’ estimates treat coincident
but distinct PAs as a single generalized area and thus
overlook differences amongst coincident PAs
(Appendix S1).
We distinguished corridors as active (n =10), planned
(n =14) or undergoing upgrades (n =9) and assessed
them at two scales: core road and/or rail infrastructure,
and a broader zone of influence defined by a 50-km-wide
swath centred on each corridor. The swaths indicate the
potential general extent of secondary effects on African
PAs on the basis of pan-tropical observations of environ-
mental change nearby roadways, including agricultural
conversion in Amazonia (~10 km distant) (Barber et al.,
2014), illegal logging in Southeast Asia (~20–30 km)
(Linkie et al., 2014) and poaching in Africa (~80 km)
(Wilkie et al., 2000; Blake et al., 2008). Rough dirt roads
or dilapidated colonial highways may occasionally exist
where planned or upgrade corridors are proposed, so that
their potential impacts would reflect enhanced traffic flows
and land-use change rather than ‘forest penetration’ per
se. Planned corridors are not certain to be developed but
are considered here in order to indicate the implications of
their development.
We estimated the conservation importance of intersected
PAs by their ‘irreplaceability scores’ for all mammal, bird
and amphibian species combined and for the threatened
species of these taxa (Le Saout et al., 2013). Scores reflect
both the number of species in a given PA and the degree to
which these species depend on the PA. They are based on
the percentage overlap between species’ global distribu-
tions and PAs. Higher irreplaceability scores are assigned
to PAs hosting larger numbers of species with larger
percentages of their distribution within the PA. For simple
guidance, a score of 1.0 is equivalent to one species being
entirely confined to a given PA, but can also be obtained if
multiple species have smaller fractions of their ranges
within the PA. Thus, a PA’s score is heavily influenced by
the extent to which it hosts species with relatively confined
distributions. Scores are nondenominational so to aid
interpretation, the scores reported here are also expressed
as proportions of the corresponding 90th percentile score
for all African PAs. The 90th percentile value was selected
as the upper value for rescaling because the maximum
irreplaceability scores for Africa are many orders of
magnitude greater and thus arguably outliers. Irreplace-
ability scores were available for 4,671 African PAs and
©2016 John Wiley & Sons Ltd, Afr. J. Ecol.
2Sean Sloan et al.
Table 1 Intersections between protected areas and african development corridors, by protected-area designation and corridor status
Protected Area Designation
Intersections by development corridors Intersections by development corridor zones
All African PAsAll corridors
Planned and upgrade
corridors All corridor zones
Planned and upgrade corridor
zones
PA count % PAs Km PA count % PAs Km PA count % PAs Km
2
PA count % PAs Km
2
PA count % PAs
Key designations 69 20.0 2,239 58 24.6 1,709 129 8.0 117,683 110 9.7 94,780 526 8.0
World heritage site 9 2.6 355 7 3.0 224 13 0.8 18,645 10 0.9 12,627 46 0.7
Ramsar site 9 2.6 527 9 3.8 422 19 1.2 22,893 17 1.5 20,489 108 1.6
UNESCO-MAB reserve 6 1.7 182 6 2.5 182 7 0.4 10,465 7 0.6 10,968 25 0.4
National park 45 13.0 1,175 36 15.3 881 90 5.6 65,680 76 6.7 50,696 347 5.3
National-grouped designations 276 80.0 5,102 178 75.4 3,138 1488 92.0 236,121 1022 90.3 162,411 6,020 92.0
Forest, nature and state
reserves
184 53.3 2,320 119 50.4 1,438 1247 77.1 102,020 855 75.5 70,863
Other protected areas and
sanctuaries
22 6.4 338 19 8.1 295 70 4.3 17,049 61 5.4 15,666
Game reserves, hunting,
and wildlife management
40 11.6 1,630 23 9.7 1,048 99 6.1 82,605 60 5.3 60,260
Community and communal
reserves
20 5.8 645 16 6.8 345 42 2.6 28,270 34 3.0 14,925
Marine national reserves,
parks and protected areas
1 0.3 11 1 0.4 11 7 0.4 965 6 0.5 646
Recreation and Safari 6 1.7 118 0 0.0 0 11 0.7 5,201 1 0.1 52
Not reported 3 0.9 41 0 0.0 0 12 0.7 12 5 0.4 0 79 1.2
Total
a
345 100 7,340 236 100 4,847 1,617 100 353,804 1,132 100 257,192 6,546 100
Total (aggregate)
b
6,115 3,948 294,536 205,983
See Table S1 for disaggregated data for national PA designations.
a
Total lengths, areas and counts of intersection are sums of intersections for the individual PA designations listed in the table. Occasional spatial overlaps between PAs of distinct
designations and a corridor/corridor zone means that these totals include partial redundancies reflecting PAs occupying the same space (Appendix S1).
b
Aggregate total lengths and areas of PA–corridor intersections overlook overlap amongst distinct PAs by ‘dissolving’ their boundaries into generalized protected areas
(Appendix S1).
©2016 John Wiley & Sons Ltd, Afr. J. Ecol.
Development corridors and protected areas 3
unavailable for 62 PAs intersected by existing or planned
corridors.
Results
Development corridors that are planned or undergoing
upgrades have particularly frequent intersections with key
African PAs. Overall, the 33 corridors intersect 69 key
PAs, but the large majority of these (58) intersect with the
23 corridors that are planned or being upgraded (Table 1).
The number of key PAs intersected by planned or upgrade
corridors is significantly greater than expected given the
number of individual corridors of each status (G
adj
=7.67,
P=0.0056; G-test for goodness of fit with Williams’
correction for sample size; Appendix S1). The three
international PA designations (WHS, Ramsar, UNESCO-
MAB) account for 40% of the key PA intersections with
corridors that are planned or undergoing upgrades
(Table 1).
The potential PA intersection length of planned corri-
dors is also considerably greater than that of upgrade or
active corridors. Planned corridors would intersect
3,612 km of PAs and have greater total, mean and
median intersection lengths of any corridor status
(Table 2). Further, of the total length of planned corridors
(19,943 km), the proportion intersecting PAs (18%) is
greater than for active (12%) or upgrade (9%) corridors
(Table 2). The lengths of PA intersections by the corridors
of each status differed significantly from expected lengths
scaled according to the total length of each corridor status
(G
adj
=515, P<0.001). Similar post hoc comparisons of
observed and expected lengths confirmed the significant
extensiveness of planned-corridor intersections specifically
(G
adj
=418, P<0.001; Appendix S1), again highlighting
their relative prominence.
Development corridors, and particularly those that are
planned or undergoing upgrades, disproportionately inter-
sect key PAs (Figure 1). Key PAs are intersected signifi-
cantly more often than expected given their frequency of
occurrence across Africa (G
adj
=54.2, P<0.0001 for all
corridors; G
adj
=63.2, P<0.0001 for planned and upgrade
corridors). Key PAs account for 20% of PAs intersected by
all corridors and 25% of PAs intersected by planned or
upgrade corridors, compared to just 8.1% for all African
PAs (Table 1). In contrast, key PAs are intersected at the
expected rate (8%) by the broader corridor zones (Table 1).
The greater than expected intersections of key PAs by
corridors rather than by the broader, more disperse
Table 2 Intersections between Protected Areas and African Development Corridors, for each Corridor Status
Corridors Status
Total
Length (km)
Intersections with corridors Intersections with corridor zones
PA count Km Mean Km Median Km PA count Km
2
Mean Km
2
Median Km
2
Planned 19,943 154 3,612 23.5 11.9 609 182,252 299.3 28.7
Upgrade 13,629 82 1,235 15.1 8.2 523 75,069 144.5 8.7
Active 19,654 127 2,493 19.6 7.8 681 128,765 189.1 16.8
Total 53,226 363
a
7,340 20.2 9.7 1813
a
386,086 212.9 16.6
Total (aggregate)
b
––6,115 28.8 7.4 –294,536 216.4 8.1
a
Counts above pertain to intersections between a PA and corridors of a given status. Where two or more corridors of different statuses intersect a single PA, that PA is counted
once for each status. This occurs rarely for corridors but regularly for broader corridor zones. Table 1 provides true counts of intersected PAs.
b
Aggregate total lengths and areas of PA–corridor intersections overlook overlap amongst distinct PAs by ‘dissolving’ their boundaries into generalized protected areas
(Appendix S1).
©2016 John Wiley & Sons Ltd, Afr. J. Ecol.
4Sean Sloan et al.
corridor zones underscores the fact that core corridor road
and rail infrastructure constitutes the epicentres of poten-
tial impact on key PAs. Two regional epicentres of
intersection between key PAs and planned or upgrade
corridors are apparent: the Western Central Congo Basin,
particularly along the iron belt of southern Cameroon and
northern Gabon and The Congo, which is driven largely by
mineral resource development (Figure 1b), and the Great
Lakes Region extending into the Eastern Savannahs
(Figure 1c), where infrastructure and mineral develop-
ment are both driving forces.
Planned and upgrade corridors also intersect PAs of
relatively high conservation value. PA-irreplaceability
scores significantly differ amongst planned, upgrade and
active corridors (ANOVA; all species F
2,280
=6.2,
P<0.05; threatened species F
2,280
=6.5, P<0.05).
Planned and upgrade corridors each have twice the
median score (~0.022; 30% of 90th percentile) of active
corridors for all species, and planned corridors alone have
twice the median score (0.001; 17% of 90th percentile) of
other corridors for threatened species.
Discussion
The scale and distribution of PA–corridor intersections
highlight apparent challenges to maintaining the
Fig 1 Protected Areas Intersected by
Development Corridors in sub-Saharan
Africa, by Protected-Area Designation
and Corridor Status. Note: Apparently
homogenous PAs of a single colour may
in fact be composed of multiple contiguous
PAs
©2016 John Wiley & Sons Ltd, Afr. J. Ecol.
Development corridors and protected areas 5
ecological integrity of key PAs. Planned corridors and
corridors undergoing upgrade to a lesser degree would
intersect key PAs of relatively high conservation value
with significantly disproportionate frequency and exten-
siveness (Table 1, Table 2). While potential impacts might
be reduced by re-routing corridors, effective conservation
will occasionally demand more substantial revisions (Lau-
rance et al., 2015a). For instance, 30% of all PA intersec-
tions are <4.2 km –a length amenable to re-routing –
although this rises to a less amenable 9.0 km for key PAs.
Further, many detours would have to be greater to buffer
PAs from any ‘halo effects’ around corridors, such as
poaching (Wilkie et al., 2000; Walsh et al., 2003; Barber
et al., 2014). Notably extensive intersections by planned
corridors include the Malagarasi-Muyovozi Ramsar wet-
lands (314 km intersected), Dja WHS (74 km), Sangha
Tri-National WHS (41 km), Niokolo-Koba WHS (40 km)
and Mole National Park (86 km).
Economic development, such as that encouraged by
development corridors, is sorely needed in Africa and is
held by some as a means of lessening human pressures
on ecologically intact hinterlands (Masters et al., 2013).
Improved spatial planning is essential to balance eco-
nomic and environmental imperatives (Wilkie et al.,
2000; Laurance et al., 2014). For example, a road
circumnavigating Tanzania’s Serengeti WHS could inte-
grate a greater human population with economic oppor-
tunities and services than a proposed ‘paved commercial
route’ that would bisect the Serengeti (Hopcraft et al.,
2015a). Nonetheless, the Tanzanian government remains
committed to the latter, imperilling one of the world’s
greatest animal migrations (Dobson et al., 2010, Africa
Geographic, 2014). In certain circumstances, careful
planning around a corridor could also enhance PA
protection, as where PA surveillance is highly irregular.
For economic development or improved surveillance to
translate into enhanced conservation depends critically on
many dynamics beyond access, however. Foremost
amongst these are whether corridors actually enhance
the economic prospects of pastoralists, farmers and others
who (over)exploit PAs’ resources (Fyumagwa et al.,
2013), whether corridors would even diminish such
exploitation, and whether concomitant increases in other
deleterious activities such as poaching or land conversion
would undo related conservation gains, with such factors
weighing more or less depending on whether one’s
perspective is short term or long term. Opinion is divided,
and there are few large-scale precedents to indicate the
likely outcome (Fyumagwa et al., 2013; Hopcraft et al.,
2015b). The status quo of African PA conservation is
wanting, and its deficiencies require resolution for
enhanced access to generally enhance PA conservation
more than it challenges it. Growing pressures on WHSs
and other flagship PAs (Osipova et al., 2014, WWF,
2015, 2016) underscore the urgency of improved spatial
planning and enhanced efforts to honour existing con-
servation commitments.
References
Africa Geographic (2014) The Serengeti Highway Battle Won, The
War with the Courts Continues. Series: Conservation, Events,
News, Wildlife. Africa Geographic. Available at: http://
africageographic.com/blog/the-serengeti-highway-battle-won-
the-war-with-the-courts-continue/ (Accessed December 2015).
Barber, C.P., Cochrane, M.A., Souza, C.M. Jr & Laurance, W.F.
(2014) Roads, deforestation, and the mitigating effect of
protected areas in the Amazon. Biol. Conserv.,177, 203–209.
Blake, S., Strindberg, S., Boudjan, P., Makombo, C., Bila-Isia, I.,
Ilambu, O., Grossmann, F., Bene-Bene, L., De Semboli, B., Mbenzo,
V. & AL, E. (2007) Forest elephant crisis in the Congo Basin.
PLoS Biol. 5.
Blake, S., Deem, S.L., Strindberg, S., Maisels, F., Momont, L., Isia, I.-
B., Douglas-Hamilton, I., Karesh, W.B. & Kock, M.D. (2008)
Roadless Wilderness Area Determines Forest Elephant
Movements in the Congo Basin. PLoS One 3, e3546.
Deguignet, M., Juffe-Bignoli, D., Harrison, J., Macsharry, B.,
Burgess, N. & Kingston, N. (2014) 2014 United Nations List of
Protected Areas. UNEP-WCMC, Cambridge, U.K.
Dobson, A.P., Borner, M., Sinclair, A.R.E., Hudson, P.J., Anderson,
T.M., Bigurube, G., Davenport, T.B.B., Deutsch, J., Durant, S.M.
& Estes, R.D. (2010) Road will ruin Serengeti. Nature 467, 272–
273.
Dudley, N. (2008) Guidelines for Applying Protected Area
Management Categories. IUCN, Gland, Switzerland.
Dur
an, A.P., Rauch, J. & Gaston, K.J. (2013) Global spatial
coincidence between protected areas and metal mining
activities. Biol. Conserv. 160, 272–278.
Edwards, D.P., Sloan, S., Weng, L., Sayer, J., Dirks, P. & Laurance,
W.F. (2014) Mining and the African environment. Conserv. Lett.
7, 302–311.
Forman, R.T.T. & Alexander, L.E. (1998) Roads and their major
ecological effects. Annu. Rev. Ecol. Syst. 29, 207–231.
Freitas, S.R., Hawbaker, T.J. & Metzger, J.P. (2010) Effects of
roads, topography, and land use on forest cover dynamics in the
Brazilian Atlantic Forest. For. Ecol. Manage. 259, 410–417.
Fyumagwa, R., Gereta, E., Hassan, S., Kideghesho, J.R., Kohi, E.M.,
Keyyu, J., Magige, F., Mfunda, I.M., Mwakatobe, A., Ntalwila, J.,
Nyahongo, J.W., Runyoro, V. & Røskaft, E. (2013) Roads as a
threat to the Serengeti ecosystem. Conserv. Biol. 27, 1122–
1125.
©2016 John Wiley & Sons Ltd, Afr. J. Ecol.
6Sean Sloan et al.
Hopcraft, J.G.C., Bigurube, G., Lembeli, J.D. & Borner, M. (2015a)
Balancing conservation with national development: a socio-
economic case study of the alternatives to the Serengeti Road.
PLoS One 10, e0130577.
Hopcraft, J.G.C., Mduma, S.A.R., Borner, M., Bigurube, G., Kijazi,
A., Haydon, D.T., Wakilema, W., Rentsch, D., Sinclair, A.R.E.,
Dobson, A. & Lembeli, J.D. (2015b) Conservation and economic
benefits of a road around the Serengeti. Conserv. Biol. 29, 932–
936.
IUCN & UNEP-WCMC (2015) The World Database on Protected
Areas. UNEP-WCMC, November 2015. Available at:
www.protectedplanet.net.
Laurance, W.F., Goosem, M. & Laurance, S.G. (2009) Impacts of
roads and linear clearings on tropical forests. Trends Ecol. Evol.
24, 659–669.
Laurance, W.F., Clements, G.R., Sloan, S., O’Connell, C.S.,
Mueller, N.D., Goosem, M., Venter, O., Edwards, D.P., Phalan,
B., Balmford, A., van der Ree, R. & Arrea, I.B. (2014) A global
strategy for road building. Nature 513, 229–232.
Laurance, W.F., Peletier-Jellema, A., Geenen, B., Koster, H.,
Verweij, P., van Dijck, P., Lovejoy, T.E., Schleicher, J. & van Kuijk,
M. (2015a) Reducing the global environmental impacts of rapid
infrastructure expansion. Curr. Biol. 25, R259–R262.
Laurance, W.F., Sloan, S., Weng, L. & Sayer, J. (2015b) Estimating
the environmental costs of Africa’s massive ‘development
corridors’. Curr. Biol. 25, 3202–3208.
Le Saout, S., Hoffmann, M., Shi, Y., Hughes, A., Bernard, C.,
Brooks, T.M., Bertzky, B., Butchart, S.H.M., Stuart, S.N.,
Badman, T. & Rodrigues, A.S.L. (2013) Protected Areas and
Effective Biodiversity Conservation. Science 342, 803–805.
Linkie, M., Sloan, S., Kasia, R., Kiswayadi, D. & Azmi, W. (2014)
Breaking the vicious circle of illegal logging in Indonesia.
Conserv. Biol. 28, 1023–1033.
Maisels, F., Strindberg, S., Blake, S., Wittemyer, G., Hart, J.,
Williamson, E.A., Aba’a, R., Abitsi, G., Ambahe, R.D., Amsini, F.,
Bakabana, P.C., Hicks, T.C., Bayogo, R.E., Bechem, M., Beyers,
R.L., Bezangoye, A.N., Boundja, P., Bout, N., Akou, M.E., Bene,
L.B., Fosso, B., Greengrass, E., Grossmann, F., Ikamba-Nkulu, C.,
Ilambu, O., Inogwabini, B.-I., Iyenguet, F., Kiminou, F.,
Kokangoye, M., Kujirakwinja, D., Latour, S., Liengola, I.,
Mackaya, Q., Madidi, J., Madzoke, B., Makoumbou, C., Malanda,
G.-A., Malonga, R., Mbani, O., Mbendzo, V.A., Ambassa, E.,
Ekinde, A., Mihindou, Y., Morgan, B.J., Motsaba, P., Moukala, G.,
Mounguengui, A., Mowawa, B.S., Ndzai, C., Nixon, S., Nkumu, P.,
Nzolani, F., Pintea, L., Plumptre, A., Rainey, H., De Semboli, B.B.,
Serckx, A., Stokes, E., Turkalo, A., Vanleeuwe, H., Vosper, A. &
Warren, Y. (2013) Devastating decline of forest elephants in
central Africa. PLoS ONE,8, e59469.
Masters, W.A., Djurfeldt, A.A., de Haan, C., Hazell, P., Jayne, T.,
Jirstr€
om, M. & Reardon, T. (2013) Urbanization and farm size in
Asia and Africa: implications for food security and agricultural
research. Global Food Security 2, 156–165.
Osipova, E., Shi, Y., Kormos, C., Shadie, P., Zwahlen, C. & Badman,
T. (2014) IUCN World Heritage Outlook 2014. IUCN, Gland,
Switzerland.
Perz, S.G. (2014) Sustainable development: the promise and perils
of roads. Nature 513, 178–179.
Walsh, P.D., Abernethy, K.A., Bermejo, M., Beyers, R., De
Wachter, P., Akou, M.E., Huijbregts, B., Mambounga, D.I.,
Toham, A.K., Kilbourn, A.M., Lahm, S.A., Latour, S., Maisels, F.,
Mbina, C., Mihindou, Y., Ndong Obiang, S., Effa, E.N., Starkey,
M.P., Telfer, P., Thibault, M., Tutin, C.E.G., White, L.J.T. &
Wilkie, D.S. (2003) Catastrophic ape decline in western
equatorial Africa. Nature,422, 611–614.
Weng, L., Boedhihartono, A.K., Dirks, P.H., Dixon, J., Lubis, M.I. &
Sayer, J.A. (2013) Mineral industries, growth corridors and
agricultural development in Africa. Global Food Security 2, 195–
202.
Wilkie, D., Shaw, E., Rotberg, F., Morelli, G. & Auzel, P. (2000)
Roads, development, and conservation in the Congo Basin.
Conserv. Biol. 14, 1614–1622.
WWF (2015) A Global Assessment of Extractive Activity within
World Heritage Sites.World Wildlife Fund for Nature.
WWF (2016) Protecting People Through Nature: Natural World
Heritage Sites as Drivers of Sustainable Development. World
Wildlife Fund for Nature, Gland, Switzerland.
(Manuscript accepted 5 September 2016)
doi: 10.1111/aje.12377
Supporting information
Additional Supporting Information may be found in the
online version of this article:
Table S1 Disaggregated Data on Intersections between
Protected Areas and Development Corridors, by Protected-
Area Designation, Corridor Type, and Corridor Status.
Appendix S1 Supplementary Text.
©2016 John Wiley & Sons Ltd, Afr. J. Ecol.
Development corridors and protected areas 7