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Conservation planning for 's Albertine Rift: conserving a biodiverse in the face of multiple threats

A NDREW J. PLUMPTRE,SAM A YEBARE,DEO K UJIRAKWINJA and D AN S EGAN

Abstract The Albertine Rift is one of Africa’s most biodi- Introduction verse , but is threatened by habitat loss as a result ‘ ’ of agricultural expansion and human development. Pre- frica has been described as the last frontier for devel- vious studies estimated that % of the region has been Aopment and there are plans to increase foreign invest-  lost to agricultural conversion and we estimate here that ment in the (Ohwoso et al., ). Most of this % is allocated for mining concessions. For conservation global interest is focused on natural resource extraction, sup- planning, we used niche models for species endemic to the ported by major infrastructure development projects planned  Albertine Rift and those that are globally threatened. We as- across Africa (Laurance et al., a). These projects include a  sessed where to conserve these species using three scenarios: growing network of roads (Laurance et al., ), pipelines,  () a baseline assuming equal conservation costs across all railways and hydroelectric dams (Laurance et al., b) grid cells in the study area, () a scenario locking in existing planned for development. Similarly, mining concessions for protected areas (i.e. always selecting them by default) and both exploration and production cover large parts of Africa, assessing which unprotected areas require conservation, and as companies in countries such as China and India (Tull,   () a scenario considering mining planned across the region. ;Kaplinskyetal., ) seek mineral resources for their Marxan analyses produced similar results for the three industries. Mining concessions are being established across scenarios, highlighting the importance of existing pro- the continent, with little thought for environmental impacts. tected areas and the value of several community-managed Although investment in Africa can help overcome poverty, or provincial protected areas in the eastern Democratic it needs to also protect the environmental services that people Republic of the Congo. The current protected area network rely upon and the biodiversity that underpins tourism as a   covers , km and an additional , km would be major source of income in many countries. Methods to min- required to ensure the conservation of all threatened and imize the impacts of development exist and mostly promote   endemic species outside the parks and wildlife reserves. the mitigation hierarchy (BBOP ;Saenzetal., ). However, if trying to avoid mining concessions this in- The first component of the hierarchy encourages avoidance  creases to , km , an area larger than the existing pro- of sites with high biodiversity and critical habitats for threat- tected areas. Some mining concessions harbour species with ened species, and is the most effective way to minimize  a restricted range and would thus need to be protected to impacts (Phalan et al., ). Conservation planning can ensure the persistence of threatened and endemic fauna help identify areas to be avoided and locations for poten-  and flora. These mining concessions should be challenged tial offsetting activities (Saenz et al., ). We show here by the conservation community. how such approaches can be used to minimize the impacts of planned mining in one of Africa’s most biodiverse Keywords Albertine Rift, biodiversity, conservation plan- regions. ning, Marxan, mining, niche models, protected areas Africa’s Western Rift Valley, the Albertine Rift, has been  Supplementary material for this article is available at designated an endemic area (Stattersfield et al., ), a   doi.org/./S Global Priority Ecoregion (Olson & Dinerstein, ; Burgess et al., ) and part of the Eastern Hotspot (Brooks et al., ; Plumptre et al., ). The richness of vertebrate and plant taxa has been documented previously (Plumptre et al., , ) and in  there were , plant species and , terrestrial vertebrate spe- ANDREW J. PLUMPTRE* (Corresponding author, orcid.org/0000-0002-9333- 4047), SAM AYEBARE,DEO KUJIRAKWINJA and DAN SEGAN Wildlife Conservation cies reported for the region. More thorough surveys, iden- Society, 2300 Southern Boulevard, Bronx, New York City, NY 10460, USA tification of new species and additional records for plants E-mail [email protected] have increased these numbers to , plants and , ter- *Also at: Conservation Science Group, Department of Zoology, University restrial vertebrates (Plumptre et al., ), and more species of Cambridge, Cambridge, UK, and Key Biodiversity Areas Secretariat, c/o BirdLife International, Cambridge, UK are being discovered and described, particularly amphibians, Received  October . Revision requested  December . reptiles and plants. This region has the highest diversity of Accepted  February . First published online  January . terrestrial vertebrates in Africa, containing . % and %

This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0/), which permits unrestricted re-use, Downloaded fromdistribution, https://www.cambridge.org/core and reproduction in any medium,. IP address: provided 170.106.35.93 the original, work on 29 is Sep properly 2021 cited. at 02:40:31, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/termsOryx, 2021, 55(2),. https://doi.org/10.1017/S0030605319000218 302–310 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000218 Conservation planning for the Albertine Rift 303

of Africa’s bird and mammal species, respectively. It also harbours more endemic and globally threatened vertebrates than any other part of the continent (Plumptre et al., ). At least  species of mammals, , reptiles, amphi- bians, fish, butterflies, dragonflies and plants are endemic to the Albertine Rift. Of the terrestrial vertebrates and plants  species are categorized as Critically Endangered,  as Endangered and  as Vulnerable on the IUCN Red List (Plumptre et al., ). Many taxa have not been fully as- sessed or are categorized as Data Deficient (poorly known and potentially threatened because they are rare and with a limited distribution; Bland et al., ), so the true number of threatened species may be higher. Protected areas in the region include four World Heritage Sites (, Bwindi Impenetrable, Kahuzi Biega and Virunga National Parks), a Man and the Biosphere Reserve (Queen Elizabeth National Park) and three Ramsar sites (Lake George and Edward wetlands, Virunga Park, and the delta of at Murchison Falls National Park; Fig. ). There is a strong positive correlation between biodiver- sity and human population density across Africa (Burgess et al., ). The Albertine Rift contains some of the conti- nent’s highest human population densities (Burgess et al., ; Cordeiro et al., ), with up to , people per  km in south-west and (Plumptre et al.,  ). As a result, the biodiversity is under threat from FIG. 1 Protected areas in the Albertine Rift region where we habitat loss and degradation, and from hunting. An esti- assessed the optimum areas for conservation, showing extent of mated % of the Albertine Rift has been converted to agriculture and mining concessions. farmland or settlement (Ayebare et al., ) as people take advantage of the rich volcanic soils around the rift valley. Recent mining developments for minerals, oil and gas are However, the strategic framework plan did not assess the further increasing the threats to biodiversity in the region critical sites that would be needed to conserve all endemic (Johnson, ; Plumptre et al., a). Governments in and threatened species in the Albertine Rift region because the five countries of the Albertine Rift have established min- data to make this assessment were not available at that time. ing concessions across much of their land, most of which are Since  the Wildlife Conservation Society and partners currently in the exploration phase. In several countries these have surveyed many areas of remaining natural habitat to exploration areas overlap with protected areas, and in the support more rigorous conservation planning across the eastern Democratic Republic of the Congo (DRC) the ex- region. traction of minerals has fuelled armed conflict and insecur- Conservation planning approaches provide methods to ity (Maystadt et al., ). assess where to focus conservation efforts to ensure persis- During – a process to develop a strategic frame- tence of all species, while minimizing the area that needs work plan for the Albertine Rift was implemented (ARCOS, to be set aside or the cost associated with these efforts ), involving international and national NGOs, univer- (Moilanen et al., ). The programmes Zonation sities, local communities and protected area authorities. (Moilanen & Kujala, ), Marxan (Possingham et al., This process identified six key landscapes of conservation ; Ball et al., ) and C-Plan (Pressey et al., ) importance that would protect many of the endemic and are commonly used for such planning. These methods threatened species: () the Murchison-Semliki Landscape, aim to efficiently invest conservation resources by either western Uganda, () the Greater Virunga Landscape, strad- identifying the minimum cost incurred to achieve a suite dling the borders of south-west Uganda, northern Rwanda of conservation targets or maximize return on a given in- and eastern DRC, () the Maiko-Itombwe Landscape, east- vestment. Planners use these tools by initially setting target ern DRC, () the Congo-Nile Divide Landscape, straddling amounts for each of the conservation features (in our study, the border of southern Rwanda and northern , species), and then dividing the landscape into planning () the Greater Mahale Landscape, western , and units within which the species composition, abundance () the Marungu-Kabobo Landscape, south-eastern DRC. or contribution towards the conservation targets can be

Oryx, 2021, 55(2), 302–310 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000218 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.93, on 29 Sep 2021 at 02:40:31, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319000218 304 A. J. Plumptre et al.

calculated. Once the cost of achieving the target in each models relating to climate (), topology (), hydrology (), planning unit has been calculated, the analysis aims to iden- geology () and human activities () that were thought like- tify the more frequently selected cells (i.e. those with unique ly to influence species distributions (Ayebare et al., ; assemblages of species) and other planning units required to Supplementary Table ). These were all raster maps of   ensure sufficient area is conserved for each cell. High quality km cell resolution. We clipped all predictor variables to species distribution data are required for these analyses. the area of interest and obtained a pairwise Pearson cor- Here, we use these planning methods to assess where to tar- relation between predictor variables using ENMTOOLs get conservation investment across the Albertine Rift, to en- (Warren et al., ). To minimize the effect of multi- sure its threatened and endemic species will survive in the collinearity and overfitting, we retained only variables with long term despite the threats from agriculture and mining , . correlation. concessions. We tested model accuracy using the area under the re- ceiver operating characteristic curve test statistic (Fielding & Bell, ; Freeman & Moisen, ) and selected model Methods outputs with values $ . for the final analysis (Manel et al.,   We modelled species ranges for  endemic and  globally ). We used % of the occurrence records for training  threatened species using the methods reported in detail in and % for testing. After assessing model accuracy, we Ayebare et al. (). We summarize the methods here. fitted the final models for all species using all occurrence records. To convert the predicted habitat suitability from a continuous logistic output format into a binary (presence/ Biodiversity data absence) output, we used the maximum training sensitivity plus specificity threshold rule (Liu et al., ), which mini- We compiled georeferenced biodiversity data from sur- mizes the mean error rate for positive observations and the veys carried out across the Albertine Rift by the Wildlife error rate for negative observations (Freeman & Moisen, – Conservation Society during . In addition, we ). obtained species distribution data for certain species from the Global Biodiversity Information Facility (GBIF, ), Chicago Field Museum (Chicago, USA), Tanzania Loss of wildlife habitat to agriculture Mammal Atlas (Tanzania Wildlife Research Institute, Arusha, Tanzania), University of Texas at El Paso (El The predicted species ranges included areas in which nat- Paso, USA), Centre de Recherche en Sciences Naturelles ural habitats had been replaced by agriculture, plantations  de Lwiro (Lwiro, DRC), and Missouri Botanical Garden or settlements. Using a combination of the GlobCover  (St Louis, USA) to provide additional georeferenced records land cover map (Arino et al., ) and maps of different for the endemic and threatened (according to the IUCN Red landscapes produced by the Wildlife Conservation Society List; IUCN, ) species of the Albertine Rift. We used a from satellite images, we estimated the extent of human- total of , presence records in the modelling process: modified areas across the Albertine Rift and clipped species  birds (,), large mammals (,), small mammals (,), ranges to exclude these areas (Ayebare et al., ). plants (,), reptiles () and amphibians (). Mapping of mining concessions

Modelling species distributions To create maps of mining concessions we used mining con- We used a maximum entropy species distribution approach cession layers from MOXI mining cadastre maps (MOXI,  to model areas of suitable habitat (Maxent ..e; Phillips ), digitizing the mining concessions from images as the et al., ). We selected Maxent because it requires only data were not available to download as a shapefile. We - species’ presence data and environmental predictor vari- tained further information on mining concessions from con- ables and performs as well or better than other species dis- tacts within government ministries in the various countries. tribution modelling techniques (Elith et al., , Phillips  et al., ). We used Maxent default parameters (auto fea- Scenarios for conservation planning tures, convergence threshold of ., maximum number of background points = ,, regularization multiplier = ) Marxan (Possingham et al., ) uses simulated anneal- to fit most of the models. However, one-third of the spe- ing to estimate the near-optimal solution to a conservation  cies (– records) were fitted using hinge features, which planning problem. We overlaid  km hexagonal cells across  are functions for piecewise linear splines and fit models the entire Albertine Rift extent, covering , km .We closely related to generalized additive models (Elith et al., assigned a cell to a species’ modelled range if . % of the ). We selected  potential predictor variables for the cell was predicted to contain the species. Target amounts

Oryx, 2021, 55(2), 302–310 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000218 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.93, on 29 Sep 2021 at 02:40:31, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319000218 Conservation planning for the Albertine Rift 305

for each species were set in Marxan, aiming to ensure the Pm is the proportion of the cell within a mining concession conservation of a viable population for each species. For and Pp is the proportion within a protected area. Applying species that are relatively abundant where they occur we this formula means that if a cell contained no mining con- selected % of the species’ current range within the cession or protected area, conserving it cost  units; con- Albertine Rift extent as a target. Some species such as ele- serving a cell entirely within a mining concession and no phants, apes and large carnivores require large areas of protected area cost  units; a cell entirely within a protected habitat to maintain viable populations. For these species, or area and no mining concession cost  unit; and conserving those that occur at low density or within a restricted area, a cell that was entirely within a mining concession and a we selected higher percentages of cells (–%) to ensure protected area cost  units. This effectively lowers the cost their needs would be met in the final conservation plan. of conservation in protected areas and increases the cost in We analysed the data for all species, and for () only glob- mining concessions, whether these occur within or outside ally threatened species (endemic and non-endemic; IUCN, protected areas. ), and () only endemic species. Marxan requires a boundary length modifier value that can be used to minim- ize boundary length and thereby identify results that are Results more compact (lower boundary:area ratio). Higher values We included a total of  endemic and globally threatened of this modifier generally result in larger areas and more species as conservation targets in the Marxan analyses, with expensive solutions. We set the boundary length modifier ranges modelled for  endemic and  globally threat-   value at . to apply minimal constraint as we were inter- ened (not endemic) species, and  endemic species with ested in the least costly solutions. only point locations in cells. The latter species were incorpo-   We ran Marxan times to obtain a selection frequency rated in the Marxan analysis as presences within the  km for each cell in the Albertine Rift extent. We analysed three grid cells, and because there were few points (, ) for each scenarios: the first scenario assumed the cost of conserva- species, we set a target of %intheMarxan analyses. tion to be equal in all cells and thus identified the minimum area required to conserve all species. This is useful as a baseline to compare with the results of other scenarios. Where to conserve if costs are equal in all cells The second scenario locked in existing national parks and reserves, meaning these areas were always selected by de- This scenario assumes conservation costs are equal for all fault. Proposed community-managed or existing provincial cells and provides a baseline assessment that shows where reserves were not locked into the analysis because these all threatened and endemic terrestrial vertebrates and plants are legally less secure and we wanted to assess their impor- would be conserved at minimum cost. Even with costs as- tance in the overall conservation plan. Provincial reserves sumed equal across all cells, the analysis showed that most established in –, including Itombwe Reserve of the important areas fall within the existing protected  (Kujirakwinja et al., ), Kabobo Reserve, Ngandja area network (Fig. ). Key areas identified outside protected Reserve, and community reserves between the Kahuzi areas included the highlands between Itombwe and Kabobo Biega and Maiko National Parks in eastern DRC, were not Massifs, forested areas around Tayna and Kisimba-Ikobo locked in (Plumptre et al., ). As well as always selecting Reserves in DRC, and woodlands and Sitebi Highlands east cells within existing parks and reserves, the Marxan analysis of Mahale Mountains National Park in Tanzania. There in this second scenario identified additional cells outside was little variation in results between the assessments protected areas that met the targets. This scenario is more of endemic vs globally threatened species (Supplementary  realistic than the first because it recognizes conservation in- Fig. ). vestments made to-date. The third scenario built upon the second by preferentially weighting cells within protected Where to conserve outside existing national parks and areas but also considering the development needs of the reserves affected countries, specifically aiming to avoid existing or potential mining concessions. Because some mining conces- In this scenario existing parks and reserves were locked into sions occur within protected areas we did not lock in the the analysis to identify additional areas required to conserve protected areas in this analysis. Instead, we weighted costs all threatened and endemic terrestrial vertebrates and plants according to whether a cell was within a mining concession outside the existing protected area system. The analysis or protected area using the formula: showed that the Tayna and Kisimba-Ikobo Reserves west of and the highlands of the Itombwe Massif C = 6 + (2A × P )–(A × P ) m p and ridge down to the Kabobo Massif are important areas where C is the relative conservation cost for a cell, A is the outside the existing parks and reserves in DRC (Fig. ).  area of the cell ( km , except for cells at the edge of a lake), The areas east of Mahale Mountains National Park

Oryx, 2021, 55(2), 302–310 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000218 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.93, on 29 Sep 2021 at 02:40:31, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319000218 306 A. J. Plumptre et al.

  FIG. 2 Scenario : selection frequency of  km cells assuming FIG. 3 Scenario : selection frequency of  km cells, locking in equal conservation costs across all cells, aiming to conserve all (i.e. always selecting) all cells within existing parks and reserves threatened and endemic terrestrial vertebrates and plants in the (but excluding existing or proposed community managed region. protected areas, shown with a dashed border).

(including Ntakata forest and the Sitebi Highlands) in Area requirements outside existing protected areas western Tanzania were also identified as important as they contain the Kungwe apalis Apalis argentea, African elephant We evaluated the area designated as irreplaceable (selected  Loxodonta africana and African wild dog Lycaon pictus. % of the time) that occurred within or outside protected   The results of this analysis were used to help establish the areas (Table ). Under scenario the irreplaceable area with- Itombwe, Ngandja and Kabobo Reserves as provincial in parks and reserves is expectedly high because these are reserves in eastern DRC. The results for globally threat- locked into the analysis and are always selected by default. ened vs endemic species are very similar (Supplementary The area of irreplaceable sites outside protected areas was   Fig. ). similar under scenario (equal cost) and scenario (avoid- ing mining concessions) but larger for scenario  (protected areas always selected), indicating that some species of con- Where to conserve in the face of future developments servation concern occur primarily outside protected areas. In terms of the area required for the most cost efficient This scenario aimed to balance the existence of protect- solution under the different scenarios, scenario  (conser- ed areas with the planned mining developments in the vation cost equal for all cells) required the smallest area. Albertine Rift to assess where conservation should be tar- Locking in the protected areas (scenario ) increased the geted considering both of these factors. Where cells are total area required by %, but the area that needs to be con- still selected within mining concessions it shows that these served outside protected areas is smallest in this scenario. sites are critical for certain species and mining should ideally The most cost effective solution under scenario  (avoid- not proceed here (Fig. ). Maps of mining concessions for ing mining concessions and preferentially selecting cells in  the region show that , km (%) of the area we fo- protected areas) increased the area required by %. The cused on is allocated to existing or proposed exploration percentage of the Albertine Rift extent required to conserve concessions. Patterns for threatened vs endemic species all endemic and threatened species ranged from .% were similar (Supplementary Fig. ). under scenario  to .% under scenario  (Table ).

Oryx, 2021, 55(2), 302–310 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000218 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.93, on 29 Sep 2021 at 02:40:31, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319000218 Conservation planning for the Albertine Rift 307

and existing mining concessions as the main threats to spe- cies conservation. These factors currently have the biggest impact in the Albertine Rift region because they are wide- spread and can occur even in remote forests. We could not assess artisanal mining because it is mapped incom- pletely across the Albertine Rift. Regulations for small-scale mining exist in all countries in the region but are often not enforced. However, given the lower impact of such mining activities (except for the hunting of some large mammal species by miners for food) they are probably less harmful to most endemic and threatened species in the Albertine Rift than industrial mining. Gorillas and elephants are most at risk from artisanal mining because they are hunted for both meat and trophies (Plumptre et al., b). Our analyses showed that it is possible to conserve all endemic and threatened species in the Albertine Rift region while al- lowing for most of the potential large-scale future mining developments. However, taking into account the existing protected area network in the region, the cost and land re- quirements to achieve this are considerably higher than in a scenario that would allow a complete redesign of pro- tected areas, without relying on any previous designations. Although most species can be preserved outside existing mining concessions, there are some areas where conserva-  FIG. 4 Scenario : selection frequency of  km cells, with mining tion has to take place within planned mining concessions concessions being more, and existing parks and reserves less to achieve the conservation targets set in Marxan. These in- costly to conserve. clude proposed mining concessions within Kahuzi Biega National Park, Tayna Reserve, Itombwe and Kabobo Massifs Discussion and concessions around Bugoma Forest Reserve in Uganda (Fig. ). Mining concessions in these locations should be The Albertine Rift region is under pressure from agricul- contested and mining companies should avoid them as tural expansion linked to high human population densities, part of any mitigation hierarchy. and from potential developments, particularly mining con- Taking into account the existing parks and reserves in the cessions. Other developments (e.g. roads, railways, power- Albertine Rift and avoiding placing conservation efforts in lines, oil pipelines and urban expansion) are also likely in mining concessions (comparing the percentage area of the this region, but these were beyond the scope of this study. best solution in scenarios  and ), an additional .%of Here, we considered existing habitat loss to agriculture the region’s area needs to be designated for conservation

TABLE 1 Results of Marxan analyses for conservation planning in the Albertine Rift region, showing the area of irreplaceable cells (selected in % of model runs) and area of the best (most cost efficient) solution, separated by areas inside and outside existing parks and reserves, for three modelling scenarios. Scenario  assumes equal conservation costs across all grid cells, scenario  locks in (i.e. always selects) cells in existing protected areas, and scenario  avoids mining concessions and prefers existing protected areas. The best solution’s total area required for conservation and the area required outside existing protected areas are also given as per cent of the Albertine Rift extent.

Scenario 1 Scenario 2 Scenario 3 Inside protected areas Area of irreplaceable sites (km2) 10,798 134,246 45,398 Area of best solution (km2) 55,181 134,246 91,154 Outside protected areas Area of irreplaceable sites (km2) 8,504 22,744 8,581 Area of best solution (km2) 86,391 64,586 145,704 Total area of best solution (km2) 141,572 198,832 236,858 Total area of best solution (% of Albertine Rift extent) 14.26 20.02 23.85 Area of best solution outside protected areas (% of Albertine Rift extent) 8.70 6.50 14.67

Oryx, 2021, 55(2), 302–310 © The Author(s), 2020. Published by Cambridge University Press on behalf of Fauna & Flora International doi:10.1017/S0030605319000218 Downloaded from https://www.cambridge.org/core. IP address: 170.106.35.93, on 29 Sep 2021 at 02:40:31, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605319000218 308 A. J. Plumptre et al.

to preserve all threatened and endemic species when avoid- the chimpanzee Pan troglodytes but rarely consider the ing mining concessions. Existing protected areas have been other taxa which are more threatened. This area requires a established not only to conserve endemic and threatened conservation project focused on the wider biodiversity of the species, but also to provide ecosystem services such as region and should be better protected (Plumptre et al., ). tourism revenue, and substantial changes to these areas are These results show that the existing protected area net- unlikely. However, conserving all threatened and endemic work is relatively well placed and is conserving many of species and avoiding mining concessions requires an addi- the threatened and endemic species of this highly biodiverse tional .% of the Albertine Rift extent to be conserved out- region. However, an additional .% of land is needed to side protected areas, compared to a scenario that does not adequately protect all species of conservation concern. For seek to avoid mining concessions (.%vs.%; Table ). many species these additional areas may not need to be This would be economically challenging given the high designated as protected areas but could be managed with human population densities in the area. Some mining con- other effective area-based conservation measures, provided cessions are not avoided by the Marxan analysis despite in- the habitat is maintained for the species. Figure  shows creased costs, implying that these sites are critical for one or where conservation efforts should focus outside protected more species of conservation concern (Fig. ). These conces- areas (see also areas listed above). Conservation practi- sions should be contested and mining companies should tioners and the governments of the region should work avoid them. However, if mining is to go ahead the com- with local communities to ensure these sites are managed panies involved should minimize impacts on endemic and together with the existing protected areas and that mining threatened species as far as possible and offset residual concessions are not allocated there. impacts elsewhere in the Albertine Rift. Offset sites should be those outside protected areas identified as important for Acknowledgements We thank the John D. and Catherine  T. MacArthur Foundation (Grant: 97201), the U.S. Fish & Wild- the conservation of endemic and threatened species (Fig. ). life Service (Grant: F13AP00680), U.S. Agency for International Conservation sites that were not locked in as protected Development/CAFEC program (AID-660-A-13-00010-64788) and areas (community or provincial reserves) and that were Wildlife Conservation Society (WCS) for funding; Eli Greenbaum consistently selected by the analyses include the Tayna (University of Texas at El Paso), Julian Kerbis Peterhans and Reserve, Kisimba-Ikobo Reserve, Itombwe Reserve, Kabobo John Bates (Chicago Field Museum) and Charles Foley and Tim Davenport (Tanzania Mammal Atlas) for providing point location Provincial Reserve, and Ngandja Provincial Reserve in east- data; and James Watson for supporting DS and advising on analyses. ern DRC. Tayna and Kisimba Reserves are important for populations of Grauer’s cuckoo shrike Ceblepyris graueri Author contributions Study conception, fundraising, model revi- and Allanblackia kimbiliensis (Clusiaceae). Itombwe Re- sions, writing: AP; species distribution models for endemic species: serve contains several endemic or near-endemic species SA; data collection and compilation: DK; initial analyses in Marxan: (Kujirakwinja et al., ) including the Itombwe river DS. frog Phrynobatrachus asper, Itombwe golden frog Chryso- Conflict of interest None. batrachus cupreonitens, Congo bay Phodilus prigo- ginei and Itombwe nightjar Caprimulgus prigoginei. Kabobo Ethical standards This research abided by the Oryx guidelines on and Ngandja Reserves also harbour several endemic ethical standards. species including Willard’s horseshoe bat Rhinolophus wil- lardi, wild ginger Aframomum ngamikkense (Zingiberaceae; Fischer et al., ), Kabogo myosorex Myosorex kabo- goensis and Kabogo crocidura Crocidura lwiroensis (Kerbis References Peterhans et al., ). These sites have been recognized for ARCOS () A Framework for Conservation in the Albertine Rift: their conservation importance but require additional pro- –. Albertine Rift Conservation Society, Kigali, Rwanda. tection or upgrading to national reserves. ARINO, O., RAMOS PEREZ, J.J., KALOGIROU, V., BONTEMPS, S., Other sites that were consistently selected outside pro- DEFOURNY,P.&VAN BOGAERT,E.() Global Land Cover tected areas included the Sitebi Highlands east of Mahale Map for  (GlobCover ). doi.pangaea.de/./    Mountains National Park in western Tanzania, the escarp- PANGAEA. [accessed October ]. AYEBARE, S., PLUMPTRE, A.J., KUJIRAKWINJA,D.&SEGAN,D. ment between Itombwe Reserve and Ngandja Reserve, and () Conservation of the endemic species of the Albertine the forest between Tayna Reserve and Maiko National Park Rift under future climate change. Biological Conservation, in DRC. Although many of these sites will protect some , –. species that also occur in the neighbouring reserves, the BALL, I.R., POSSINGHAM, H.P. & WATTS,M.() Marxan and Sitebi Highlands are particularly important as they con- relatives: software for spatial conservation prioritisation. In Spatial  Conservation Prioritisation: Quantitative Methods and tain several endemic plants, endemic butterfly species Computational Tools (eds A. Moilanen, K.A. Wilson & (Kielland, ) and the Kungwe apalis Apalis argentea H.P. Possingham), pp. –. Oxford University Press, (Moreau, ). Projects in this region tend to focus on Oxford, UK.

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