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Plumptre, A.J., T.R.B. Davenport, M. Behangana, R. Kityo, G. Eilu, P. Ssegawa, C. Ewango, D. Meirte PDF

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BIOLOGICAL CONSERVATION 134 (2007) 178–194 available at www.sciencedirect.com journal homepage: www.elsevier.com/locate/biocon The biodiversity of the Albertine Rift Andrew J. Plumptrea,*, Tim R.B. Davenportb, Mathias Behanganac, Robert Kityoc, Gerald Eiluc, Paul Ssegawac, Corneille Ewangoa, Danny Meirted, Charles Kahindoc, Marc Herremansd,k, Julian Kerbis Peterhanse,f, John D. Pilgrimg,l, Malcolm Wilsonh, Marc Languyi, David Moyerj aWildlifeConservationSociety,P.O.Box7487,Kampala,Uganda bWildlifeConservationSociety,P.O.Box1475,Mbeya,Tanzania cMakerereUniversity,P.O.Box7062,Kampala,Uganda dRoyalMuseumforCentralAfricaatTervuren,Leuvensesteenweg11,3080Tervuren,Belgium eRooseveltUniversity,UniversityCollege,430SMichiganAvenue,Chicago,IL60605,USA fFieldMuseumofNaturalHistory,Chicago,1400S.LakeShoreDrive,Chicago,IL60605-2496,USA gCenterforAppliedBiodiversityScience,ConservationInternational,1919MStreetNW,Suite600,Washington,DC20036,USA hP.O.Box178,Groblersdal0470,Mpumalanga,SouthAfrica iWWFEasternAfricaRegionalProgrammeOffice(EARPO),P.O.Box62440,00200Nairobi,Kenya jWildlifeConservationSociety,P.O.Box936,Iringa,Tanzania kNatuurpunt.Studie,Coxiestraat11,2800Mechelen,Belgium lBirdLifeInternationalinIndochina,4/209DoiCan,BaDinh,Hanoi,Vietnam A R T I C L E I N F O A B S T R A C T Articlehistory: TheAlbertineRiftisoneofthemostimportantregionsforconservationinAfrica.Itcon- Received16June2005 tainsmorevertebratespeciesthananyotherregiononthecontinentandcontainsmore Receivedinrevisedform endemicspeciesofvertebratethananyotherregiononmainlandAfrica.Thispapercom- 17July2006 piles all currently known species distribution information for plants, endemic butterfly Accepted8August2006 speciesandfourvertebratetaxafromtheAlbertineRift.Theliteratureonfishspeciesrich- Availableonline14November2006 nessandendemismisalsoreviewedtoassesstheimportanceofthelargerlakesintheRift forconservation.Weusedatafrom38protectedandunprotectedareastoprioritisesites Keywords: withintheAlbertineRiftforconservationbasedupontheirnumbersofendemicandglob- AlbertineRift allythreatenedspecies.VirungaandKahuziBiegaNationalParksandItombweMassifin Prioritysetting DemocraticRepublicofCongo,BwindiImpenetrableandKibaleNationalParksinUganda, Mammals andNyungweNationalParkinRwandarankhighestintermsofnumbersofbothendemic Birds and globally threatened species. Six conservation landscapes are described that include Reptiles mostofthesesitesanditisarguedthatafocusontheselandscapesmaybeamoreholistic Amphibians methodtoensurethesafetyofthepriorityareasoftheAlbertineRift. Plants (cid:2)2006ElsevierLtd.Allrightsreserved. Biodiversity 1. Introduction etal.,2003).ThispartofAfricacontainsthe‘Mountainsofthe Moon’orRwenzoriMassifthatincludesAfrica’sthirdhighest The Albertine Rift is the most species rich region for verte- peak, the Virunga Volcanoes made famous by its mountain bratesontheAfricancontinent(Brooksetal.,2001;Plumptre gorillas, active volcanoes in the Virunga National Park, and * Correspondingauthor. E-mailaddress:[email protected](A.J.Plumptre). 0006-3207/$-seefrontmatter (cid:2)2006ElsevierLtd.Allrightsreserved. doi:10.1016/j.biocon.2006.08.021 BIOLOGICAL CONSERVATION 134 (2007) 178–194 179 Lake Tanganyika – Africa’s deepest lake. The Albertine Rift FosseyGorillaFundInternational,InstituteforTropicalForest encompassesmuchofthewesternRiftvalleydowntosouth- Conservation,InternationalGorillaConservationProgramme, ernTanzaniaandnorthernZambia.Wedefinetheregionas MakerereUniversityInstituteofEnvironmentandNaturalRe- extending from 30km north of Lake Albert to the southern sources,WildlifeConservationSociety(WCS)andWWF.The tip of Lake Tanganyika, including the valley, flanks of the compilation of the biodiversity information was one aspect escarpment and associated protected areas, and the range ofthisplanningprocessandwasledbyWCS.Thispaperad- ofspeciesendemictoit(Plumptreetal.,2003).Variousother vancesthis collaborative effortbyusingthecollateddatato publicationsalsorecognizethe‘AlbertineRift’(Poulsen,1997; prioritisesitesintheAlbertineRiftforconservationaccording Prigogine,1985):itisanEndemicBirdAreaaccordingtoBird- totheirnumbersofendemicandgloballythreatenedspecies. Life International (Stattersfield et al., 1998), WWF have de- fined it as a ‘Global-200’ priority ecoregion (Olson and 1.1. TheAlbertineRiftregion Dinerstein,1998;Burgessetal.,2004),andConservationInter- national(CI)hasrecognizeditaspartoftheEasternAfromon- As part of this planning process a series of meetings were tane Hotspot in their second global analysis (Brooks et al., heldbetweenFebruary2001andMarch2003,whentheframe- 2004).ThereismuchoverlapinthedefinitionoftheAlbertine workwasfinallydeveloped.Duringthisprocessitwasagreed Riftbetweenthesepublications,butalsosomedifferencesin thatthedefinitionoftheAlbertineRiftshouldbeasinclusive geographicalcoverageaseachanalysishasuseddifferentcri- aspossibleforthemomentsothatovertimeitcouldbere- teria to delineate the region. WWF’s Ecoregion focuses pri- fined (Plumptre et al., 2003). The current adopted definition marily on the montane forests and separates moorland thereforeincludesallthenaturalhabitatswithin100kmeast from these forests, while BirdLife International’s Endemic oftheborderofDemocraticRepublicofCongo(DRC)andfol- birdareaisbroaderandincludesthemoorlandsandhighland lowsthe900mcontourlineineasternDRC,includingthepro- swamps. CI’s Eastern Afromontane Hotspot is broader still tected areas in northern Zambia (Fig. 1). The 900m contour and includes the definition of the Albertine Rift used here was selected because there are museum collections at the (seebelow). RoyalMuseumforCentralAfricainTervuren,Belgium,ofAlb- TheAlbertineRiftisnotonlyimportantforitsbiodiversity ertineRiftendemicbirdspeciesthatwerefoundaslowasthis butalsoforitsecologicalprocessesandecosystemservices. altitude. Thesavannaparkscontainedsomeofthehighestbiomasses The total area encompassed is around 313,000km2 oflargemammalsrecordedonearthinthe1960s(Cornetd’Elz- (Plumptre et al., 2003). The habitats range from the glaciers ius,1996).Warandpoachinghaveledtomajordecreasesinthe and rock at the top of the Rwenzori mountains (5100m), numbersoflargemammalsintheseparksbutmostofthespe- down through alpine moorland (3400–4500m), Giant Senecio ciesarestillpresentandcouldrecovertoformerlevelswith good protection (Plumptre et al., 2007). The impacts of the browsingandgrazingoftheelephants,hippopotamuses,buf- falos,andantelopespecieshadamajorinfluenceonthevege- tationoftheparks(Delvingt,1978)andasaresultofthedecline innumbersofthesespeciesitisthoughttheparksarechanging (Eltringham,1999).ThevolcanoesintheVirungaNationalPark areactiveandinfluencetheecologyofalargeportionofthis parkanditssurroundings.Thefisheriesinsomeofthelakes arethemostproductiveonthecontinentandprovidealiveli- hoodformanypeople(Beadle,1974;Snoeks,2000)andtheriv- ers and streams flowing from the forests on the mountains providecleanwater.InRwanda,forexample,itisestimated thatmorethan70%ofpeopleobtainwaterthatcomesfrom theirnationalparks(Weber,1989).Thespectacularlandforma- tionsandrichbiodiversityoftheAlbertineRiftmeanthatithas greatpotentialfortourism.Civilwarsandinternationalcon- flictoverthepast30yearshavehamperedtourismdevelop- mentbutwhenpeacecomestotheregionthereisenormous potentialtodevelopworldclasstourism. Thispapersummarisestheexistingliteratureonbiodiver- sitysurveyswithinsitesintheAlbertineRift.Itistheresultof a collaborative effort between many different NGOs, pro- tectedareaauthoritiesand museumsthathaveinformation about this region. Since 2001 the protected area authorities and their NGO partners have been developing a strategic frameworkforconservationintheAlbertineRift.Atthistime Fig.1–MapoftheextentoftheAlbertineRiftasconsidered acoreplanninggroupwasestablished,whichdevelopedthe here.Forestedprotectedareas(orsurveyedareas)arein strategic framework for conservation in the Albertine Rift, darkgrey,savanna/miombowoodlandprotectedareasare that includes the Albertine Rift Conservation Society, Dian inlightergrey. 180 BIOLOGICAL CONSERVATION 134 (2007) 178–194 and Lobelia vegetation (3100–3600m), giant heather (3000– et al. (1987). A list for the Virunga Volcanoes was produced 3500m), raised bogs (3000–4000m), bamboo forest (2500– using de Witte (1938), Wilson (1995) and Hutterer et al. 3000m), montane forest (1500–2500m), to lowland forest (1987).ForDRC,deWitte(1938)reportedonextensivesurveys (600–1500m), savanna woodland (600–2500m) and savanna by Belgian scientists in the Virunga National Park. Muhlen- grassland(600–2500m).PapyrusandCarexwetlands,together bergetal.(undated)providedalistforKahuziBiegaNational withlakesandstreams,havetheirownuniquehabitattypes ParkandOmarietal.(1999)providedalistforItombweMas- varying from the rocky and sandy edges to the benthic and sif. For Burundi, a listwasobtainedforKibiraNational Park bathypelagic zones in the depths of the lakes. Several very (FieldMuseum,Chicago(KerbisPeterhans,inlitt.,FMNHdata- specialised habitats also occur as a result of the volcanic base,Chicago),PeaceCorpsandINECN(undated).ForTanza- activity in the Virunga National Park, including lava flows nia,listswereobtainedforGombefromtheGombeNational and their associated colonising vegetation, hot springs and Parkwebsite,forMahalefromAnonymous(1985)andforMbizi speciesadaptedtocarbonmonoxideandmethane. fromD.MoyerandW.Stanley’ssurveys.ForZambia,listsfor Sumbu and Mweru-Wantipa National Parks were generated 2. Methods fromthedistributionrecordsgivenbyAnsell(1978). 2.1. BiodiversityoftheAlbertineRift 2.1.2. Birds ThemainsourceforUgandawasthechecklistofbirdsofthe InitiallyWCScompiledlistsofspeciesfromexistingliterature national parks (Wilson, 1995) and the biological surveys and by working with researchers at Makerere University, undertaken by the Uganda Forest Department (Howard and Uganda. Much of the literature was in unpublished reports Davenport,1996–whichalsocompileddatafrommanypre- or‘greyliterature’.Dataonmammals,birds,reptiles,amphib- viously published sources). Additional data came from the ians, butterflies and plants were compiled, and information Enhancement of Research Capacity (ENRECA) project man- on some museum specimens collated with the help of mu- aged by Makerere University Institute of Environment and seum experts (Royal Museum for Central Africa, Tervuren, NaturalResources,surveysundertakenbyWCS,andrecords Belgium; Missouri Botanical Gardens, USA; Field Museum, compiled by Malcolm Wilson. Additional published records Chicago, USA, and Royal BotanicGardens,Kew, UK)from 38 thatwereincorporatedafterexpertpeerreviewincludeEvans siteswithintheAlbertineRift(Fig.2).Wealsoreviewedthelit- and Balmford (1992), Gnoske and Marks (1997), Kalina and erature for fish species totals as it was recognized that the Butynski (1996), Friedmann and Williams (1970), Dehn and lakesintheAlbertineRiftregionarealsoimportantforcon- Christiansen(2001), Stubblefield(1993), and Allan (1994).For servation. We attempted to compile information from as Rwanda,KunkelandKunkel(1969),Dowsettetal.(undated), manyareas oftheAlbertineRiftaspossible.Dataon collec- Dowsett (1990) and Plumptre et al. (2002) produced lists of tions of amphibians and reptiles from the Royal Museum birdsfortheNyungweForestReserve.Thesedatawerecom- forCentralAfricaallowedsomeanalysistobemadeofspe- bined with unpublished sightings from the WCS project in ciesoutsideprotectedareas.Datafromplantfamilydescrip- this forest. A list for the Virunga Volcanoes was produced tionsandherbariawerealsousedtogenerateapreliminary using Schouteden (1938), Wilson (1982), and Wilson (1995). list of endemic plant species, many of which are not found Schouteden (1938) reported on extensive surveys by Belgian inthesitesselectedhere.Notsurprisingly,however,mostof scientistsintheVirungaNationalParkinDRCandVerheyen thesitesselectedwereprotectedareasasthatiswheremost (1947)addedspeciestothenorthernpartofthepark.M.Lan- surveyshavebeenmadeintheAlbertineRift.However,some guyprovidedadditionalobservations.Muhlenbergetal.(un- unprotectedareasalsohadsurveyinformation,includingthe dated) and Wilson and Catsis (1990) provided a preliminary ItombweMassif,MarunguMassif,highaltitudeareaswestof list for Kahuzi Biega National Park and thiswas augmented LakeEdwardandMtKaboboaswellasareasaroundMahale by M. Herremans using the database of bird specimens at MountainsNationalPark.WhilenotcoveringthewholeAlber- theRoyalMuseumforCentralAfricainTervuren.Schouteden tineRift,thesesitesdoincludethelargerprotectedareasand (1949) surveyed Katanga district from which a list for Mar- thelargestareasofunprotectedhabitat. unguMassifwasderived,Prigogine (1960)providedalistfor Themainsourcesusedforeachtaxonwereasfollows: Mt Kabobo, Prigogine (1971–1984), Wilson and Catsis (1990) and Omari et al. (1999) provided lists for Itombwe Massif 2.1.1. Mammals and Prigogine also provided lists for Idjwi Island (Prigogine, ThemainsourceforUgandawasthechecklistofmammalsof 1967)andfortheareawestofLakeEdward(Prigogine,1953). thenationalparks(Wilson,1995)andthesmallmammalsur- ManynewrecordsineasternDRChavebeenpublishedbyDe- veys undertaken by the Uganda Forest Department(Howard mey et al. (2000). Tom Butynski contributed records for Mt and Davenport, 1996). Additional data for large mammals Tshiaberimu. Schouteden (1966) published a list of birds of camefromKingdon(1971–1983)andforsmallmammalsfrom Burundi with locations where they had been sighted – this Kerbis Peterhans et al. (1996), Kerbis Peterhans and Austin publication was used to compile a list for Ruzizi National (1996),KerbisPeterhans(1997)andvanderStraetenandKer- Park.Gaugrisetal.(1981)addedspeciestothislistandINECN bisPeterhans(1999).Dowsett(1990)producedalistofmam- producedalistforKibiraNationalParkinthemid1980s(IN- mals for the Nyungwe Forest Reserve in Rwanda and this ECN, undated). van de Weghe and Loiselle (1987) also pro- wascombinedwithunpublishedsightingsfromtheWildlife duced a list for Bururi forest reserve. Neil Baker provided ConservationSocietyprojectinthisforestandfromHutterer listsforGombeStreamandMahaleMountainsNationalParks BIOLOGICAL CONSERVATION 134 (2007) 178–194 181 Fig.2–Thenorthern(a),central(b)andsouthern(c)portionsoftheAlbertineRiftshowingthelocationsofthevarious protectedareas(NP=nationalpark;FR=forestreserve;WR=wildlifereserve)orungazettedareaswithspeciesdata(no suffix).Darkershadedareasareforestedandlighterareasaresavannagrasslandorwoodland. from the Tanzania Bird Atlas database. These were used to getheralistforKibaleNationalPark.Spawlsetal.(2002)iden- correct and add to lists compiled by Stanford and Msuya tify localities for species for East Africa using general maps (1995), Ulfstrand and Lamprey (1960), and Moreau (1943). D. and these were used to assign species to a site if this was MoyerprovidedlistsforMbiziforest. either mentioned in the text or if the map distribution was unequivocal. M. Behangana, from recent surveys, provided 2.1.3. Reptiles severalrecords.HinkelandFischer(1988)wasusedtodevelop Pitman(1938,1974)wasusedasastartingpointforreptilesin alistofspeciesforVirungavolcanoesandNyungweforestin Uganda.DrewesandVindum(1998)providedaspecieslistfor Rwanda.ThiswasaugmentedbydeWitte(1941)fortheViru- BwindiImpenetrableNationalParkandVonesh(1998)putto- nga volcanoes. Dowsett (1990) also provided a list for 182 BIOLOGICAL CONSERVATION 134 (2007) 178–194 Nyungwe Forest. D. Meirte extracted lists of specimen loca- Maramagambo forests. Synnott (1985) provided a checklist tionsfromthedatabaseattheRoyalMuseumforCentralAfri- ofplantsforBudongoForestReserve.ForRwanda,plantspe- ca in Tervuren. For DRC, de Witte (1941) produced a list of cieslistsfortheVirungaVolcanoeswereobtainedfromBurtt reptiles for Virunga National Park and mapped the distribu- (1934),Robyns(1948–1955),Troupin(1978–1988),andtheher- tionsofchameleonsincentralAfrica(deWitte,1965).D.Mei- barium at the Karisoke Research Station. Plant species for rteextractedlistsofspecimenlocationsfromthedatabaseat Nyungwe were compiled from Robyns (1948–1955), Troupin theRoyalMuseumofCentralAfricainTervuren.Onlylimited (1992),Plumptreetal.(2002)andtheherbariumattheProject datawereobtainedforBurundi.Spawlsetal.(2002)wasused Conservation de la Foreˆt de Nyungwe. Robyns (1948–1955) tocompilealistofreptilesforGombeandMahaleMountains providedarelativelycompletelistforVirungaNationalPark ParksinTanzania. inDRCandFischer(1996)providedalistforKahuziBiegaNa- tionalPark.NoplantlistswereobtainedforsitesinBurundi. 2.1.4. Amphibians ForTanzania,thelistofplantsofGombewasprovidedbyRoy DrewesandVindum(1994,1998)andDrewesetal.(1992)pro- Gereaufromhissurveysthere.ToshisadaNishidakindlypro- vided a species list for Bwindi Impenetrable National Park videdalistofplantseatenbychimpanzeesforMahaleMoun- andVonesh(1998)puttogetheralistforKibaleNationalPark tains National Park. Additional species for Mahale were inUganda.ManyrecordswereprovidedbyM.Behanganafrom obtained from Vollesen and Bidgood (1996, 1999). A species recentsurveys.InRwanda,HinkelandFischer(1988)wasused listforMbiziforestwascompiledfromMwasumbi(2000).Leb- todevelopalistofspeciesforVirungavolcanoesandNyungwe runandStork(1991–1997)wasusedtocorrectsynonymstoa forest.ThiswasaugmentedbydeWitte(1941)fortheVirunga standardisedlistofnames. volcanoes.Dowsett(1990)alsoprovidedalistforNyungweFor- est.ForDRC,deWitte(1941)producedalistofamphibiansfor 2.1.7. Datacleaningandidentificationofspeciesof VirungaNationalParkbuttheidentificationsneededupdating. conservationconcern ThedatabaseattheAfricaMuseuminTervurenisinthepro- Aftersixmonthsofdevelopingdraftlistsameetingwascon- cessofbeingupdatedforamphibianstoreflectcurrenttaxon- vened by the core planning group to bring together experts omyandweusedthecorrecteddatainthisdatabasetocreate fromtheregion,particularlyfrommuseumsandotherinsti- lists of species for DRC and to add records to sites. Laurent tutions to review and augment the draft species lists. This (1972)increasedtheamphibianlistofdeWitteforVirungaNa- meeting also addressed threats to these sites and refined tional Park and corrected some mis-identifications. Laurent the listsof endemicspeciesforthe AlbertineRift. Following (1964)wasusedtocompilealistofamphibiansfortheItombwe thismeetingtherewasanotherperiodofsixmonthsadding Massif.D.Meirteprovidedcorrectionstothetaxonomyofthe tothedataandinparticularcheckingspeciesnamesandsyn- olderliteratureandalsoaddedmanyspeciesfromthedatabase onymsbecausetaxonomyofmanyspecieshaschangedover attheRoyalMuseumforCentralAfricaatTervuren.Somedata the70yearsofpublicationsthatwereused.WCSandMake- ondistributionandsystematicsoftreefrogsweretakenfrom rereUniversitywereinvolvedinthisdatacleaning. Schiøtz (1999). Only limited data were obtained for Burundi Numbersofendemicandgloballythreatenedspecieswere andnoneforTanzania. calculatedforeachsiteandusedtoprioritisesitesintheAlb- ertineRiftforconservationas follows.Endemicspecieslists 2.1.5. Butterflies were compiled with the help of experts in each taxonomic InformationwasdrawnfromavarietyofsourcesincludingCar- field.ThemammallistcamefromConservationInternational casson(1961,1975),d’Abrera(1980,1997),Henning(1988),Kiel- (J.Pilgrim)withadditionsbyJ.KerbisPeterhans.Thelistcom- land(1990),Larsen(1991),Ackeryetal.(1995),Davenport(1996), piledbyStattersfieldetal.(1998)wasusedforbirds.Thispub- Howard and Davenport (1996), Congdon and Collins (1999), licationrecognizedtwoareasofendemisminthisregion;the Congdonetal.(2001),aswellasnumerousworkersfromearlier Albertine Rift and the Eastern Zairean Region of endemism. partsofthelastcenturyreferencedinthesepublications.Addi- However,somespeciesendemictotheAlbertineRiftoverlap tional information came from collections held at Makerere substantiallywiththosefromtheEasternZaireanregion(M. UniversityZoologyMuseum,Kampala,andtheNationalMuse- Herremans–datafromAfricaMuseumatTervuren)andthe umsofKenya,Nairobi.SteveCollins(ABRI,Nairobi)provided planninggroupmadethedecisionthatitwasbettertocom- considerable and invaluable information, and we are very binethesetwoareasinthisanalysis.Listsofendemicreptile grateful to C. Congdon (Tanzania) and A. Gardiner (Zambia) andamphibianspecieswerecompiledbyD.Meirte.T.Daven- forveryusefulcommentsonanearlierdraft. portcompiledalistofendemicbutterflyspeciesincollabora- tionwithS.Collins,C.CongdonandA.Gardiner.E.Ndomba,P. 2.1.6. Plants Ssegawa,G.EiluandA.Plumptreworkedwithbotanistsatthe The main starting point for Uganda was the tree surveys RoyalBotanicGardens,Kew,andMissouriBotanicalGardens undertaken by the Uganda Forest Department (Howard and todevelopapreliminarylistofendemicplantspeciesbased Davenport,1996).AdditionaldatawasaddedbyG.Eilu(climb- uponpublishedfamilydescriptionsfrommostexistingAfri- ers)Poulsen(1997,terrestrialherbs)andD.Hafashimana(epi- canfloras(EastAfrica,Congo,Cameroon,Gabon,andZambe- phytes) for several forests. WCS has also been surveying siaca).Thislistislikelytomissmanyspeciesbecausemany many of the Rift forests over the past year and the species familieshavenotbeendescribedinDRCorEastAfrica.Ende- identified were incorporated in the database by D. Nkuutu. mic species lists are given in Plumptre et al. (2003) and are Lock(1977)providedalistofspeciesforQueenElizabethNa- also at www.albertinerift.org. Globally threatened species tional Park. Nabanyumya (1991) listed trees for Kalinzu and were derived from the 2002 IUCN Red List (Hilton-Taylor, BIOLOGICAL CONSERVATION 134 (2007) 178–194 183 2000andupdatedlistsontheassociatedwebsitewww.iucn- maximumrank(4inthecaseofreptiles).Ameanrankwasthen redlist.org)andBirdLifeInternational(2000). calculatedforbothendemicandgloballythreatenedspecies foreachsiteacrossthefivetaxabysummingranksfromeach 2.2. Rankingsitesforconservation taxonanddividingthisbythenumberoftaxaforwhichthere weresurveydata(i.e.,forwhichaninitialrankwasevenpossi- While there is a danger that ranking sites in terms of their ble).Thesefinalstandardisedscoreswerethenranked(1–38)to conservation value can lead to the lower-ranked sites being rankalltheavailablesitesforwhichwecompileddata.Thus, sidelined, it is still important that priorities are identified thefinalrankingsallowbettercomparisonofsiteswithvarying given the limited human and financial resources available levelsofknowledge.Inpresentingtheresultswegroupedsites for conservation. However, even the poorest sites for which intocategoriesofhigh,mediumandlowrankratherthanusing we have compiled data are rich, in a global context, have actualrankingvaluesforeachsite.Thisavoidsfalseprecision manyrestricted-rangespecies,andthereforedeserveconser- becausethedataformanysitesareincomplete,somemean vationattention. rankingscoresarebaseduponjustonetaxon,andtheeffort Ranking using species data requires subjective decisions, madeinsamplingsitesvarieswidely.Comparingtherankings particularlywhendataareincomplete.Onlyfivetaxa(mam- forendemicandgloballythreatenedspeciesallowsananalysis mals,birds,reptiles,amphibiansandplants)canbeusedinthis oftherelativeimportanceofthesetwocriteria. studyoftheRiftbecausespeciesdataarenotcollatedoravail- The ranking process described above did not attempt to ableforothertaxa.Intheabsenceofmoreextensivedata,we correctfortheareaofthesites.Weweremoreinterestedin canonlyhopethatthesetaxaactasgoodsurrogatesforothers. which are the priority sites for species conservation in the Severalaspectsofsurrogacyweretestedbetweenthesetaxa. AlbertineRiftratherthanwhicharetherichersitesrelatively Thisindicatedthat,inmanycases,onetaxoncouldactasarea- when area is standardised. From a conservation perspective sonablesurrogatefortheotherfouracrossthesitessurveyed the larger a site is, the easier it is to manage compared to (Plumptreetal.,2003).Numbersofendemicspeciesforonetax- manysmallersitesofthesame totalarea; conservingafew oncorrelatedwell(PearsonCorrelations)withnumbersofen- large sites will in general be the cheaper option. However, demicspeciesinanothertaxon,forallpossiblecombinations forcomparison with other studieswe did make this correc- (P<0.001foralltests).Surrogacyforthreatenedspecieswas tion.Wecorrectedforareabycalculating not so good. Numbers of threatened mammals, amphibians scoredendemicorthreatenedspecies andbirdscorrelatedwellbetweensitesbutthreatenedplants ¼lnðnumberof speciesþ1Þ=lnðareaÞ: andreptilesdidnotshowanysignificantcorrelationwithother taxa.However,plantsandreptileshavebeenlesscompletely assessedforthreatenedspeciesthantheotherthreetaxa.Only 3. Results tworeptileswereclassifiedasthreatenedinthisregioninthe 2002RedList.Iftotalnumbersofspeciesareusedtoranksites Theresultsshowthatthisregioncontainsmorethanhalfof thenplantswilldominatetherankingsbecauseofthelarger continentalAfrica’sbirdspeciesandnearly40%ofitsmammal numberofspecies.Thegeneralpublicmaywishtoranksites species.Reptileandamphibianspeciesdonotappearasabun- onmammalandbirdfaunabecausethesearepopular.Alterna- dant but this may be a function of the effort that has been tively,itmightbebettertoprioritisesitesonspeciesthathave madeincollecting,identifyingandcataloguingtheminthisre- economicvalueandcanattracttouristsorprovidelivelihoodto gion.Whileinvertebratetaxahavebeenpoorlysurveyed,this localcommunities,orbytheirecologicalroleintheecosystem. regionisknowntohavealargenumberofendemicbutterflies Weightingscouldbemadeforcertainspeciesorcertaintaxato (probably the best surveyed invertebrate taxon). The results incorporatetheseideas.Herewehavedecidedtoweighttaxa presentedherearethecurrentstateofknowledge,whichwill equallysothatahighnumberofmammalspeciesatasitere- beimprovedoninduecourseasmoresurveysareconducted. ceivesthesamerankasahighnumberofplantsirrespective ofthe actualnumberofspecies.This effectivelygives equal weighttoanytaxonandranksbiodiversityperseratherthanso- 3.1. Biodiversityvalues cial,economicorecologicalfunctionsofbiodiversity. Siteswererankedintermsofthenumberofendemicand 3.1.1. Mammals numberofgloballythreatenedspeciesforeachtaxon.These NoendemicfamiliesoccurintheAlbertineRiftbuttwoende- rankingscoreswerestandardizedforeachsitebydividingby micgeneraoccur,Rwenzorisorex,andDelanymys.Atotalof402 themaximumrankscoretoaccountforthefactthatsomesites mammalspecies (158 generaand 46 families)have been re- do not havedata forall taxa and to account for thevarying cordedintheAlbertineRift,ofwhich35areendemic(Table1). numberofendemicorthreatenedspeciesbetweentaxa.For Most of the endemic mammals are shrews and rodents. Of example,onlytworeptilespeciesarecurrentlylistedasthrea- thelargerendemicmammals,theeasterngorillaisthebest tenedofthe175speciesfoundintheAlbertineRiftsites(pri- knownandhastwosubspecies(mountain,Gorillaberingeiberin- marily because reptiles have not been well assessed). Three gei,andGrauer’s,Gorillaberingeigraueri,gorillas).TheRwenzori siteshavethreatenedreptiles,whichmeansthatallsiteswith duiker,Cephalophusrubidus,andthegoldenmonkey,Cercopithe- no threatened reptiles scored a value of ‘4’ in the rankings. cus kandti, are two other medium sized endemic mammals. Howeverarankvalueof‘4’forbirdsisahighvaluewithmany Smallmammalshavebeenpoorlysurveyedthroughoutmuch threatened species and therefore ranks cannot be summed oftheRift,particularlytowardsthesouthernend,anditisvery across taxa without standardizing them by dividing by the likelymorespecieswouldbeaddedwithfurthereffort. 184 BIOLOGICAL CONSERVATION 134 (2007) 178–194 Table1–RichnessofmammalsacrosssitesoftheAlbertineRiftMountains,includingnumberofspecies, numberofAlbertineRift(AR)endemicspecies,andnumberofgloballythreatenedspecies(CR=CriticallyEndangered, EN=Endangered,VU=Vulnerable) Site Speciesno. ARendemicspecies ThreatenedCR,EN,VU BudongoFRa 95 0 5 BugomaFR 38 0 4 BugunguWR 9 0 1 BururiFR 9 1 1 BwindiImpenetrableNPa 135 20 7 EchuyaFRa 24 7 1 ForestWestofLakeEdward 8 0 1 GombeNP 19 1 4 IbambaroFR 2 0 0 ItombweMassif 72 4 10 ItwaraFR 18 0 0 KagombeFR 14 0 3 KahuziBiegaNPa 136 15 14 Kalinzu–MaramagamboFRa 58 1 3 KarumaWR 57 0 4 Kasyoha–KitomiFRa 47 2 3 KibaleNPa 115 5 7 KibiraNPa 71 8 7 KitechuraFR 17 0 1 KyamburaWR 37 0 3 MafugaFR 20 3 1 MahaleMountainsNPa 52 1 6 MatiriFR 12 1 0 MbiziFR 23 1 2 MurchisonFallsNPa 109 0 5 Mweru-WantipaNPa 50 0 7 NyungweNPa 86 14 3 QueenElizabethNPa 97 0 6 RwenzoriMountainsNPa 102 18 10 SemlikiNPa 86 1 5 SemlikiWR 69 0 4 SumbuNPa 61 0 6 VirungaNPa 196 21 13 Total 402 35 36 a Reasonablysurveyedforallmammals. Virunga National Park in eastern DRC has the highest Althoughthisisthemostthoroughlysurveyedgroupofani- numberofendemicmammalspecies(21).BwindiImpenetra- mals,newspeciesfortheRiftcontinuetobeaddedasmigrant bleNationalPark(20)andRwenzoriMountains(18)ranknext species and new range extensions are recorded. Within our highest. definitionoftheAlbertineRifttherearetwocontiguousEnde- Thirty-fourmammalspeciesaregloballythreatened(Crit- micBirdAreas(EBAs)definedbyBirdLifeInternational:Alber- ically Endangered, Endangered or Vulnerable) according to tineRiftandEasternZaireanLowlands(Table2)(Stattersfield the 2002 IUCN Red List (Hilton-Taylor, 2000; www.iucnred- etal.,1998).Wehavecombinedtheseareasbecauseoldmu- list.org), of which 12 are Albertine Rift endemics. These in- seumcollectionsofAlbertineRiftendemicspeciesshowthey clude eastern gorilla (Gorilla beringei), golden monkey and occuratloweraltitudesineasternDRCandthattheyoverlap Rwenzori otter shrew Micropotamogale ruwenzorii. Kahuzi inaltitudinalrangewiththeEasternZaireanlowlandspecies Biega National Park had the highest number of globally (Boberetal.,2001;Herremansetal.,2002).Thetotalnumber threatenedmammals(14)followedbyVirungaNationalPark of endemic birds in the Albertine Rift as defined here is 41 (13). species,whichincludestheendemicsofthesetwoEBAs. The Itombwe Massif, an unprotected area west of the 3.1.2. Birds northern end of Lake Tanganyika, contains more endemic Three genera are endemic to this region, Pseudocalyptomena, speciesthananyothersiteintheAlbertineRift(34).Itisclo- Graueria,andHemitesiabutnofamiliesareendemic.Atleast selyfollowedbyKahuziBiegaNationalPark(30)andVirunga 1061 bird species (in 368 genera and 80 families) occur in NationalPark(27). theAlbertineRift(Table 2) ofwhich 4.5%are migratoryspe- Twenty-fiveAlbertineRiftspeciesaregloballythreatened, ciesthatoverwinterintheregionbutdonotbreed,orwhich 13 of which are endemics. These include congo bay owl pass through on migrations within the African continent. (Phodilusprigoginei),itombwenightjar(Caprimulgusprigoginei), BIOLOGICAL CONSERVATION 134 (2007) 178–194 185 Table2–RichnessofbirdsacrosssitesoftheAlbertineRiftMountains,includingnumberofspecies,numberofAlbertine Rift(AR)endemicspeciesasdefinedbyBirdLifeInternational(withnumberofeasternZaireanlowlandendemicspeciesin parenthesesthatwereincludedinthedefinitionofAlbertineRiftEndemicspecieshere–seetext)andnumberofglobally threatenedspecies(CR=CriticallyEndangered,EN=Endangered,VU=Vulnerable) Site Speciesno. ARendemicspecies ThreatenedCR,EN,VU BudongoFR 362 0 1 BugomaFR 221 0 1 BururiFR 155 13 3 BwindiImpenetrableNP 381 24(1) 6 EchuyaFR 136 14 2 ForestsWestofLakeEdward 420 25 11 GombeNP 267 0 2 Idjwi 150 2 1 ItombweMassif 583 34(4) 15 ItwaraFR 183 0 0 KagombeFR 121 0 0 KahuziBiegaNP 335 32(3) 11 Kalinzu–MaramagamboFR 393 4 1 Kasyoha–KitomiFR 308 2 1 KibaleNP 327 3 3 KibiraNP 211 21 7 KitechuraFR 90 0 0 KyamburaWR 450 0 6 LenduPlateau 317 6 4 MafugaFR 130 10 0 MahaleMountainsNP 250 2 1 Marungu 282 1 0 MatiriFR 119 0 0 MbiziFR 116 0 0 MtKabobo 231 18 3 MurchisonFallsNP 476 0 7 NyungweNP 280 26 7 QueenElizabeth 594 0 7 RusiziNR 182 1 3 RwenzoriMountainsNP 241 21 4 SemlikiNP 441 7(5) 9 SemlikiWR 435 0 3 VirungaNP 706 27(2) 11 Total 1061 41(6) 25 IntheanalysespresentedinthispaperAlbertineRiftandEasternZaireanLowlandendemicspecieswerecombinedbecauseoftheextensive overlapindistributionsofthetwogroupsascanbeseenhere. kungwe apalis (Apalis argentea), grauer’s rush warbler (Brad- SixteenendemicreptilespeciesoccurintheRift,ofwhich ypterusgraueri)andgolden-napedweaver(Ploceusaureonucha). VirungaNationalParkcontainsthehighestnumber(11),fol- Itombwehadthehighestnumberofgloballythreatenedspe- lowedbyRwenzoriMountainsNationalPark(9)andNyungwe cies(15)followedbyVirungaNationalPark(11),KahuziBiega National Park (8). Endemic species include five chameleons National Park (11) and the mountains west of Lake Edward such as the strange-horned chameleon (Bradypodion xenorhi- (11). num), and Johnston’s chameleon (Chamaeleo johnstoni), two colubridsnakes,oneviper,sixskinks,onewormsnake(Lepto- 3.1.3. Reptiles typhlops)andonelacertidlizard. Noendemicgeneraorfamiliesofreptilesareknownfromthe Only two globally threatened reptiles are currently listed AlbertineRift. A total of 175 reptiles, from 69 genera and 20 fortheAlbertineRift(TrionyxtriunguisandOsteolaemustetra- families (about 14% of Africa’s reptiles) have been recorded spis). However, this is because the region has few data and fortheAlbertineRift(Table3).Farfewersiteshavebeensur- reptiles as a whole have not been assessed completely for veyed for reptiles to the same extent as for mammals and theirthreatenedstatus.TheIUCNGlobalReptileAssessment birds, although some records existed for at least 33 sites. isunderwayanditislikelymanymorespecieswillbeadded Itombwe Massif and Kahuzi Biega National Park in eastern totheRedList.Asaresultnositehasmorethanoneglobally DRCmaycontainmanyspeciesbuttodatehavebeenpoorly threatenedreptile(Table3). surveyed. Other areas that need work include the Marungu MassifineasternDRCatthesouthernendofLakeTanganyika 3.1.4. Amphibians andtheMahaleMountainsNationalParkanditssurrounding Three amphibian genera are endemic to the Albertine Rift; naturalvegetation. Laurentophryne, Chrysobatrachus and Callixalus. There are 119 186 BIOLOGICAL CONSERVATION 134 (2007) 178–194 Table3–RichnessofreptilesacrosssitesoftheAlbertineRiftMountains,includingnumberofspecies,number ofAlbertineRift(AR)endemicspecies,andnumberofgloballythreatenedspecies(CR=CriticallyEndangered, EN=Endangered,VU=Vulnerable) Site Speciesno. ARendemicspecies ThreatenedCR,EN,VU BudongoFR 48 1 0 BugomaFR 9 0 0 BugunguWR 9 0 0 BururiFR 1 1 0 BwindiImpenetrableNP 34 6 0 EchuyaFR 4 0 0 ForestsWestofLakeEdward 6 3 0 GombeNP 1 0 0 ItombweMassif 35 5 0 ItwaraFR 10 0 0 KahuziBiegaNP 69 7 0 Kalinzu–MaramagamboFR 9 0 0 KarumaWR 15 0 0 Kasyoha–KitomiFR 9 0 1 KibaleNP 56 3 0 KibiraNP 3 2 0 KyamburaWR 12 0 0 L.Rukwa 7 0 0 L.Tanganyika 13 0 0 LenduPlateau 6 0 0 MafugaFR 17 2 0 MahaleMountainsNP 4 0 0 MarunguMassif 6 0 0 MbiziFR 3 0 0 MtKabobo 6 2 0 MurchisonFallsNP 32 0 1 NyungweNP 43 8 0 QueenElizabeth 34 0 0 RusiziNR 3 0 0 RwenzoriMountainsNP 34 9 0 SemlikiNP 49 0 0 SemlikiWR 33 0 1 VirungaNP 109 11 0 Total 175 16 2 speciesofamphibiansintheAlbertineRift,including29gen- (Snoeks, 2000). Only Lake Malawi has more endemic fish in eraand11families,(about19%ofAfrica’samphibians). Africa. Fifty-six fish species are endemic to lakes George Thirty-sixendemicspecieshavebeenidentified.Itislikely andEdward,whileKivuandAlberthave15andsixendemic thatmoresurveyeffortwoulduncoverfurtherendemicspe- fish respectively. Only 10% of Lake Tanganyika’s shore has cies.VirungaNationalParkhadthehighestnumberofende- beenexploredandatotalofover1200faunalspecies(verte- micspecies(16)withItombweMassif(16)andNyungwePark brates and invertebrates) have been recorded, making it the (14).Fourendemicspecies(Hyperoliuspustulifer,Schoutedenella secondhighestrecordeddiversityforanylakeonearth(Patt- loveridge,Schoutedenellamossoensis,Schoutedenellavercammeni), ersonandMakin,1998). have only been recorded outside the 40 sites described here and are thus not found in any of the protected areas in the 3.1.6. Butterflies AlbertineRift. ThetotalnumberofbutterflyspeciesfoundintheAlbertine SixteenAlbertineRiftamphibiansaregloballythreatened, Riftisunknown,asthisinformationcannotbecompiledun- of which 14 are endemic. Itombwe Massif has more threa- til many more areas have been surveyed, particularly in tened species (CR, EN or VU) than other sites (11) followed eastern DRC. In Uganda, inventories of the forests in the by Virunga National Park with 10 and Bwindi Impenetrable AlbertineRifthaveshownthatatleast581speciesofbutter- NationalParkwithsix(Table4). fly,16%oftheestimated3630speciesinAfrica,occurinthis part of the Albertine Rift alone (Howard and Davenport, 3.1.5. Fish 1996). It is possible that, given the numbers from Uganda While no attempt was made to put together species lists of and Tanzania, up to 1300 butterfly species might occur in fishfortheriversandlakesintheAlbertineRift,theliterature theRift,about35%ofAfrica’stotal.Therearenoknownen- wassearchedfordataonthemajorlakes(Albert,George,Ed- demicfamilies,butthegenusKumothalesisrestrictedtothe ward,KivuandTanganyika).LakeTanganyikaalonehas289 AlbertineRift.Itisknownthat117endemicspeciesfrom49 endemicspeciesthatmakeup89%offishdiversityofthelake generaexistintheAlbertineRift(Plumptreetal.,2003).The BIOLOGICAL CONSERVATION 134 (2007) 178–194 187 Table4–RichnessofamphibiansacrosssitesoftheAlbertineRiftMountains,includingnumberofspecies,number ofAlbertineRift(AR)endemicspecies,andnumberofgloballythreatenedspecies(CR=CriticallyEndangered, EN=Endangered,VU=Vulnerable) Site Speciesno. ARendemicspecies ThreatenedCR,EN,VU BudongoFR 32 1 1 BugomaFR 20 1 0 BururiFR 4 4 1 BwindiImpenetrableNP 29 6 6 EchuyaFR 19 5 1 ForestsWestofLakeEdward 6 6 3 ItombweMassif 23 16 11 ItwaraFR 19 0 0 KahuziBiegaNP 25 7 4 Kalinzu–MaramagamboFR 25 2 2 KarumaWR 16 0 0 Kasyoha–KitomiFR 16 3 2 KibaleNP 33 5 3 KibiraNP 1 0 0 KitechuraFR 15 0 0 KyamburaWR 14 0 0 MafugaFR 1 1 0 Marungu 19 1 0 MatiriFR 15 0 0 MtKabobo 8 7 5 MurchisonFallsNP 14 0 0 NyungweNP 33 14 5 QueenElizabethNP 10 1 1 RwenzoriMountainsNP 25 7 1 SemlikiNP 24 1 0 SemlikiWR 13 0 0 VirungaNP 65 16 10 Total 119 36 16 total number of endemics (of which 85% are forest depen- crease greatly when lower plants and little studied growth dent) is considerably larger than the 78 endemic species forms such as climbers, epiphytes, lichens and bryophytes found in the Eastern Arc Mountains and coastal forests of are included. Western Tanzania, especially around Mahale TanzaniaandKenya,andcouldincreasewithfurthersurvey Mountains National Park, appears to be particularly rich in effort. Whether or not butterflies are indicative of other plantspeciesandyethasstillnotbeensurveyedintensively. invertebrate species is unclear but these numbers do dem- As such it deserves more attention. Virunga National Park onstrateunequivocallythatthisregionisnotonlyimportant in eastern DRC and Bwindi Impenetrable National Park in for vertebrate conservation. Uganda had the highest numbers of plant species recorded butbothsiteshavebeenrelativelyintensivelysurveyed.The 3.1.7. Plants Marungu Massif and Itombwe Massif in eastern DRC have Higherplantshavebeenrelativelywellsurveyedintheforests few records but also could be relatively rich and require of Uganda and Rwanda but elsewhere in the Albertine Rift surveys. surveyshavebeenpatchy.Currently5793plantspecies(from 1537generaand233families)havebeenrecordedwithinthe 3.2. Sitepriorityrankings Riftbutthiswillchangeassurveysarediscoveringnewspe- cies regularly even within Uganda (Table 5). These data in- Rankingsweremadefornumbersofbothendemicandglob- clude ferns and higher taxa but do not include the allythreatenedspeciesforalltaxa(mammals,birds,reptiles bryophytes and lichens, which are very poorly surveyed. amphibiansandplants),foreachsiteasexplainedinSection Thenumberofplantspeciesishighcomparedwithmanyre- 2.2(Table6).Wethengroupedthesitesintohigh(rankscores gions of similar size and forms 14% of all mainland Africa’s 1–12),medium(rankscores13–24)andlow(rankscores25–38) estimatedplantspecies.Apreliminaryestimateofthenum- scoringsitesforbothcriteriaandplottedtheresultsinatwo- bers of endemic plant species has been compiled by the waytableforendemicandgloballythreatenedspecies(Table WCS’s Albertine Rift Programme and this now numbers 551 7).Giventhegapsinthedataanddifferencesinsamplingef- species.Theselistsarebasedonpublishedfloradescriptions fortbetweensiteswebelievethatgroupingthesitesintothe ofplantfamilies,butmanyfamilieshavenotbeendescribed threebroadrankingcategoriesprovidesamoreconservative forDRCorEastAfrica.Someinputwasmadebyherbariaex- approachbyalleviatingthesebiases. perts, but the list is still incomplete. As a result this list Thosesixsitesthatscoredhighestareconsideredtobethe should be considered to be very preliminary and could in- mostimportantbecausetheyrankhighlyfornumberofboth

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within the Albertine Rift for conservation based upon their numbers of endemic and glob- The Albertine Rift is the most species rich region for verte-.
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