Africa's Hotspots of Biodiversity Redefined
Author(s): Wolfgang Küper, Jan Henning Sommer, Jon C. Lovett, Jens Mutke, Hans Peter
Linder, Henk Jaap Beentje, Renaat Sylva Angèle Rosine Van Rompaey , Cyrille Chatelain,
Marc Sosef, Wilhelm Barthlott
Reviewed work(s):
Source: Annals of the Missouri Botanical Garden, Vol. 91, No. 4 (Dec., 2004), pp. 525-535
Published by: Missouri Botanical Garden Press
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Volume
Number
2004
91
Annals
of the
Missouri
Botanical
Garden
4
AFRICAXS
HOTSPOTSOF
BIODIVERSITYREDEFINED1
@9
<'>
WolfgangKuper,2^9 Jan Henning Sommer,2
Jon C. Lovett,3Jens Mutke,2Hans Peter
Linder,4Henk Jaap Beentje,5Renaat
Sylva Angele Rosine VanRompaey,6
C^rille Chatelain, Marc Sosef,8 and
WilhelmBarthlott2
AB>TITACT
A key prol)lemforconservation
is the!( ointi(lente of regiotlsof higll})io(liver<,itw
withregionsof higllhumanimpact.
'I'ssellty-five
of tlle mostthleatenedtellter! of plant(3iVeIsitwereidentifiedb) Myer!et al., andthese ' hotspots"play
a (rutial 101ein international
(onselvation.>tategiesThe primarygoalof the hotspotsis to coverthe lllostthreatened
(ellters of plallt diversity,hut theireffi(aty 11asnot yet l)eell tested enll)irialls. 'or sul)-Saharan
Africa,our study
ezvaluates
tlle hotspotspostulatedly Myersand cozlparesthetllto a set of redetfirle(l
hotspotsproposedoll tlle basis of
Illappeddistriloution
data for 5985 plant speties. The two sets of hotspotsoverlapby 485tc Ourredefinedhotspots
include80%of the speciesancl669 ot the range-restri(
ted speciesof tlle suh-Saharan
florain areasunderhighhuman
' Data compilation,as well as the coordirlation
of the mergeddatabasesat the Nees Institutefor Biodiversityof
Plantsin Bonn,are fundedby the GermanFederalMinistryof Educationand Research(BIOLOG
BIOTAProgramme,
wwwbiota-africa
org),andthe AkadellliederWissenschaften
undderLiteratur,
Mainz,andsupportedby the University
of Bonn CABS-Conservation
International
andthe DanishCentreforTropicalBiodiversity
(Universityof Copenhagen,
Denmark)fundedthe plantcompilationworkin York,in particularfor JamesTaplin.The DanishCentreforTropical
Biodiversityand variousotherfundssupporteddatacompilationin Denmark,in particularforAnne-Marie
Burgerand
ChristianFrimodt-M011er
We thankthe numerousexpertswho contributedto the plant data used in this paper,in
particularPatriciaCraven(Windhoek),
LaurentGautier(Geneva),TonyRebelo(Kirstenbosch),
Jer8meDegreef(Meise),
PeterFrankenberg
(Stuttgart),
Don Kirkup(Kew),NorbertJurgens(Hamburg),
SigridLiede(Bayreuth),RogerPolhill
(Kew),AdjimaThiombiano
(Ouagadougou),
MauricioVelayosandFernandoCasas(Madrid),
JanWieringa(Wageningen),
and GeorgZizka (Frankfurt).
Paul Williamsof The NaturalHistoryMuseum,London,providedthe WORLDMAP
software.JanSchnitzler(Bonn)assistedwiththe manuscript.
WethankGeroldKier(Bonn),TonyRebelo(Kirstenbosch),
Paul Williams,and twoanonymousreviewersforvaluablecommentson the manuscript.
2 Nees Institute
for Biodiversityof Plants,Universityof Bonn,Meckenheimer
Allee 170, D-53115 Bonn,Germany.
[email protected].
3CentreforEcology,Lawand Policy,Environment
Department,
Universityof York,Heslington,York,YO105 DD,
UnitedKingdom.
4 Instituteof SystematicBotany,Zollikerstrasse
107, CH-8008,Zurich,Switzerland.
5 RoyalBotanicGardens,
Kew,Richmond,SurreyTW93AE, UnitedKingdom.
6 Universite
librede BruxellesULB,Laboratoire
de Botaniquesystematique
et Phytosociologie,
50 Av.F.D Roosevelt
1050 Bruxelles,Belgium.
7 Conservatoire
et Jardinbotaniquesde la Ville de Geneve,case postale60, CH-1292Chambesy,Switzerland.
8 NationalHerbarium
of the Netherlands-Wageningen
branch,BiosystematicsGroup,WageningenUniversity,GeneraalFoulkesweg37, 6703 BL Wageningen,
The Netherlands
9 Corresponding
author.
ANN.MISSOURI
BOT.GARD.91: 525-535. 2004.
Annals of the
Missouri Botanical Garden
526
impact,whereasthesevaluesare 15% and11%lowerforMyers'shotspotsDespitehavingequalsize anda considerable
spatialoverlapwithMyers'shotspots,ourredefinedhotspotsincludefurtherhighlythreatenedcentersof plalltdiversity
ill the Maputaland
PondolandRegion,in Ikatanga,the East AfricanAfromontane
region,the LowerGuineaRegion,
alld the AlbertineRift Manyof these redefinedhotspotsare poorlyprotectedcentersof plantand animaldiversity.
Theirconservation
is essentialfor a comprehensive
coverageof Africa'scentersof biodiversity.
Kez ?Xords: Africa,biodiversity,
conservation,
endemism,GlobalStrategyfor PlantConservation
(GSPC),hotspots,
important
plantareas.
Thereare growingconcernsaboutthe coinciding
spatial [)attelnxof humanpopulationdensityand
biodivelsitF((3incottaet al., 2000; Myerset al.,
2000; 13alnlf'ol(l
et al., 2001; Lucket al., 2004). In
a pioneerillgstu(lyin 1988, Myersidentifieda global set of hotxots whereexceptionalconcentrations
of species with high levels of endemismface exceptionaltheats of destruction(Myers,1988). Later up(latex(IVlyel;,1990;Myerset al.,2000) played
a crll(ial lole ill the publicperceptionof the threat
to lio(liversit+alld the developmentof large-scale
ollcel)txlol settlngspatla prlorltleslorln sltUconselvation.as Ie(lLlired
by Article8 of the ConventioIIon [3iologi( al Diversity(UNCF,D,199:2).
Whereax('t'lltelS of diversityare identifie^(l
on the
|)asis of liologi( al richnessan(lell(lemism.the hotsI)ots( ol( esl)tof Myerset al. (2()()())coml)illedtwo
Clitelitl:lulblst(liversityand the threatsto thatdiversit. ll] l)la(ti(evhotspotswere(lefine(las areas
with less than '30%intactprirmary
vegetbltion
and
withat lea!it()..5Xc
of the globalplantspeeies being
endermicto tlle area.
Myels et al. based their study on two types of
infornlation.
ln additionto expertadvice,theyused
summalyinformationon the numberof endemic
species occulringin the hotspots.This "inventorybased type'!of information(Barthlottet al., 1999;
Mutkeet al., 2002) was the only availal)leinformationon the biodiversityfor manyareas (Myers
et al., 2000; Krupnik& I(ress, 2003), thoughthe
applicationand the analysis of this data type is
limited(Mutke& Barthlottin press).Indeed,Myers et al. (2000; see also Brookset al., 20()1)mentioned several areas in sub-SaharanAfrica for
which the availabledataare insufficientto decide
whetheror not they representa hotspot.Theyalso
statedthatdue to the heterogeneous
datasituation,
theyhadto use the endemismcriterioneventhough
it was felt to be "minimalist"
(Myerset al., 2000).
In addition,informationon biodiversitywas, and
oftenstill is, only availablefor comparatively
large
areas delineatedby politicalboundaries.This low
spatialresolutionlimitsthe accuracyto whichcenters of diversity can be identified (I(rupnik&
Kress,2003).
Morethan a decade afterthe first global set of
,.
.
.
.
.
.
n
.
.
hotspotswas published,it has becomepossibleto
define hotspotsmore rigorouslyand accountably.
Diversitycan now be calculatedfrom a "taxonbased type"of information(Barthlottet al., 1999;
Mutke& Barthlott,in press),consistingof reliable
data on the indivictualdistributionareasof a large
numberof taxa (Williams,1996; Burgesset al.,
1998). New high-resolutionsocioeconomicdata
(CIESIN,2000; Sandersonet al., 2002) have become available.Complexalgorithms(Margules&
Pressey,2000) allow a moreprecise identification
of centersof richllessand endemism(I,ovettet al.,
2000; Linderet al.! in press)and permitanalyses
of the degreeof potentialconflictbetweenconservation and lan(l-LIseinterests (Balmfordet al.,
2001; Willianlset al.! 2003; Lucket al.?2004).
Consequent
ly. taxon-basedbiodiversityinformation has been lSe(l to criticizethe globalhotspot
areas as define(ll)y Myerset al. (2000) for their
conceptuall)a(kglound(Jepson& Canney,2001),
for their perfortllatl(ein comparisonto otherarea
networksseleeted on more complex parameters
(Williamset cll.! 1996; Balmfordet al., 2001), for
their size (Brumtllit& Lughadha,2003), and occasionallyfor theil location(Burgesset al., 2002;
Krupnik& Krex9^2003). However,to our knowledge, there is no study that assesses whetherthe
hotspotsdo indee(tcoverthe mostthreatenedcenters of plant (livelsity,even thoughthis was the
Orlglnalntentlonot t 11S pRlOrltlZatlOn SC aeme.
In this pa)el we evaluatethe extent to which
Myers'shotspots include a maximumnumberof
rareplant species in those areas most threatened
by humanimpactin Africa.This is importantfor
four reasons:(i) as the foundationof food webs,
plants are of essential importancefor terrestrial
biodiversity,(ii) plantsmightbe the best available
surrogateto representinvertebrate
diversity(Myers,
1988);(iii) plant diversityis insufficientlycovered
by currentnetworksof protectedareas(Burgesset
al., in press);and (iv) plants are the groupfrom
whichhotspotswere identifiedby Myerset al.
Which criteriashouldbe used to evaluateand
enhancethe performance
of the hotspots?A modificationof the currenthotspotswouldbe desirable
if theyexcludeareasin whichmorerareplantspe.
.
.
.
p
.
.
.
.
.
Volume 91, Number 4
2004
Kuper et al.
Africa's Biodiversity Hotspots
527
these areasis an importantchallengeforthe future.
Inadequateavailabilityof distributiondata could
have two effects on our analysis:First, in cases
wherewe have inadequatedatafor areasincluded
in the hotspotsas delineatedby Myerset al. (2000),
testingthe hotspotson the basis of our datamight
METHODS
result in an underestimation
of their performance
in coveringsub-Saharanplant diversity.However,
SPECIES
DATABASE
in fact the rnostdetailed,updated,andcomprehenSince 2003! an internationalgroupof research sive distributiondata in our databaseis for those
institutionshas contributeddata on Africa-wide areascoveredby the Myers'shotspots(whichmay
plant distriloutions
to the BiogeographicInforma- partlybe a consequenceof highlightingthese areas
tion S)stem on AfricanPlant Diversit^(BISAP), since 1988): (i) Data for the upper Guinea(West
whichis hostedandcuratedby the BIOMAPS
Proj- Africa)hotspotstem fromthe NationalHerbarium
ect withinthe BIOLOGBIOTAframework(www. of the Netherlands-Wageningen
branch;(ii) datafor
the WesternCape are from the NationalBotanic
biota-africa.org).
The dataleaseincludes Africa-widedistribution Institute,Republicof SouthAfrica,andcontributed
recordsfor 6269 species (statusMarch2004), all by T. Rebeloand N. Jurgens;and (iii) distribution
of whichhad been taxonomicallyrevised.The da- datafor East Africastem froma compilationof retabase comprisesabout 330,000 distributionrec- stricted-rangespecies for the area of the Floraof
ordsfromconfirmedcollectiolllocalities.The spa- TropicalEastAfrica(H. Beentje unpublisheddata,
tial precisionof the datavariesfromexactlocalities togetherwith othersources).Even thoughthe data
(mainlyfromherbariumcollectionswithgeo-refer- for WestAfricaare comparativelycomprehensive,
enced localities)to one-degreeresolutiondatafrom the easternpartof the upperGuineahotspotis podigitizedmaps. Data are organizeclin MS Access tentiallyless well re)resented due to lower data
databasesand have been plottedand analyzedus- availahility.Second it is possiblethatthe selection
ing WORLDMAPsoftware(Williartls2002) ancl of ouralternativeset of hotspotson the basis of our
ArcView3.2a C71S
software(ESRI 2000). In or(ler data misses areas that are richer in species than
to achieve maximumcomparal)ility
with previous those includedsimply}-ecausethe formerones are
analysesoll sub-Saharanzoodiversity(Balmforcl
et not adequatelycollected. This is certainlya key
if areasare selectedon the basis of speeies
al.,2001; Brooks et al.^2001; Burgesset al., 2002; problern
Burgesset al.^in press; Fjeldsa et al., 2004; De richnessalone. But this problemis not as promiKlerket al., 2004), all plantdistril)ution
datawere nent when selecting hotspots,because their idenrescaled to a one-degreegrid resolutionwithina tificationis also l)ased on the intensityof human
base map of 1713 one-degreelatitude-longitude impact.Hotspotstend to be particularlywell colgrid cells coveringmainlanclsub-SaharanAfrica lected (i) due to their popularitywith biologists
southof 17 degreeslatitude.By restrictingthe geo- (Reddy & Davalos,2003) and (ii) because their
graphiccoverageto Africasouthof the Saharaand high humanimpactis associatedwith a comparaexcludingthose species foundonly on offshoreis- tively good infrastructure
providingaccess (Gibbs
lands, a databasewith 5EJU5
plantspecies remains Russell et al., 1984).
forfurtheranalyses.This is between10Wo
and 15Wo
of the species in the Africanflora(I,brlln & Stork, DIGITIZING THE HOTSPOTS OF MXTERSET AL.
1991-1997; Beentje et al., 1994). AdditionalinIn orderto comparethe sub-Saharan
hotspotsof
formationon the origin of this data set is documentedin Burgesset al. (in press,see alsofootnote Myerset al. (2000) with our dataarrangedin onedegreecells, we firstdigitizedMyers'shotspotpoly1).
Ourplantdata are the mostcomprehensive
ever gons (delineationstaken from Mittermeieret al.,
assembledfor the study area, but inevitablyhave 1999) and overlaidthem on our grid of cells. We
limitations.Thereare certainareasin Ethiopiaand then identifiedall cells that had a spatialoverlap
Somalia,Sudan,the CentralAfricanRepublic,the of morethan 25@o of their area with the hotspots
Republic of Congo, the DemocraticRepublic of polygons(see methodsin Burgesset al., 2002). The
Congo,and Angolawhereanalysisof samplingin- decisionto acceptor omitcells withless than25@c
of the hotspots
tensity indicates that plant diversityis not ade- overlapcouldaffectthe performance
quatelydocumentedin our databases(W. Kuper, in covering Africa's most threatenedcenters of
unpublisheddata). The scientific explorationof plantdiversitycomparedto alternativehotspotsdecies are morethreatenedby humallimpactthanin
included areas. Here we test the efficacy of the
hotspotsusing the most comprehensiveavailable
databaseof plant species distributionsfor sub-SaharanAfrica.
Annals of the
Missouri Botanical Garden
528
30
o
o
-
cZ
H
sx520
o
a
cZ
o
o ,,,
cZ
o
*
.N
n
O
cZ
M----
*-*
10
o
0
25
50
75
Human Footprint (%)
100
Figure1. Scatterplot
illustratingthe valuesof range-sizerarityandmeanhumanfootprint(rescaledfromSanderson
et al., 2002) for each of the 1713 sub-Saharan
one-degreecells (eachrepresentedby a blackdot,partlyconcealedby
othersymbols).Bothparameters
are rescaledto percentages.Forbettervisualization,
the sevencells witha range-size
rarityof morethan30%arenot shown(X/Y-valuesforthese cells: (63/100),(75/89),(99168),(97/60),(38142),(53136)
(42/35) these are all in bothMyershotspotsand redefinedhotspots).Opencircles:125 cells coveredby the Myers
hotspots.Grayboxes: 125 cells with highestproductof range-sizerarityand humanfootprint,cited as "redefined
hotspots"in the text.
lineatedon the basis of ourdata.Wethereforetested whetherthe inclusionof furthercells (thosewith
less than 25Wooverlap)wouldimprovethe performanceof the hotspotsin comparisonto our redefined hotspotsof equal area in each case. If this
was the case, we acceptedthe cell as partof the
hotspots.Withthis mostconservativemethodin favorof the performance
of the Myerset al. hotspots,
125 cells wereidentifiedas beingpartof thesehotspots. Fromhere on, they will be referredto as
"Myershotspots"(Fig. 2A).
CHOOSING DATA TO REPRESENT PLANT ENDEMISM
AND ITS THREAT
The aim was to select data to best approximate
the twocriteriaused for the delineationof the Myers hotspots, which were plant endemism and
threat.Using our data we selected 125 cells in
which plant diversityand threat are maximized
(Fig. 1) on the basis of the followingtwomeasures.
(i) Wecalculatedrange-sizerarityforeachcell (Fig.
2B). This combinesthe numberand rangesizes of
species in each cell (Williams,1996;Ikier& Barthlott, 2001; Wieringa& Poorter,2003). The more
species that occur in a cell and the smallertheir
rangesare, the higherthe value. We chose rangesize raritysince it best approximates
the endemism
criterionappliedby Myerset al. (2000).In contrast
to theircriterion,range-sizerarityincludesnotonly
the species strictly endemic to the hotspots,but
also everyspecies whoserangeoverlapswiththem.
Hence, two areas with the same numberof strict
endemicscan still be differentiatedaccordingto
their contributionin coveringthe rangesof other
species. (ii) As a surrogateforthreat,we calculated
the mean humanfootprintfor each cell, rescaling
the datafromSandersonet al. (2002)to one-degree
cells (followingthe methodsused in Balmfordet
al., 2001; Lucket al., 2004). Forcoastalcells, the
meanfootprintwas calculatedon the basis of mainland valuesonly.The humanfootprintindexshows
similarspatialpatternsto parameterssuch as populationdensity,whichwas used in previousstudies
(Balmfordet al., 2001; Lesslie, 2002; Luck et al.,
2004), but in additiontakes into considerationa
widerrangeof factors,such as infrastructure
and
land-cover.Thus,agricultural
areasthatdo notnecessarilyhave a high humanpopulationdensityare
includedas havinga high humanimpacton biodiversity.We calculated mean values of human
Figure2.
A. Mapof the set of redefinedhotspotsidentifiedin the presentstudy,contrastedwiththe hotspotsas
delineatedby Myerset al. (2000).Solidblackline withincontinent:Myershotspots.Grayopensquares:125 one-degree
gridscoveredby the Myershotspots(see methods).Red squares:125 cells coveredby the redefined
hotspots(cellswith
highestproductof range-sizerarityand meanhumanfootprintper cell). Blackdotsrepresent125 firstcells (compare
Table1) of a near-minimum-cost
areaset (Williams,2002, the entirenear-minimum-cost
set is illustratedin Fig. 2B).
B. Range-sizerarityper one-degreegridcell. This measurecombinesthe valuesforrichnessandthe rangesizes of
the species occurringin each cell. It is calculatedas the sum of the inverserangesizes per cell (Williams,1996).
Volume 91, Number 4
2004
Kuper et al.
Africa's Biodiversity Hotspots
529
Figure2.
B. (Continued) Notethatmanyspecies of mediumrangesize mayresultin a similarcell sTalue
as one
withfewerspecies of very smallrangesize. Blackdots mark422 cells thatformthe near-minimum-cost
areaset for
sub-Saharan
plantdiversityin the dataset (Williams,2002). Thesecells representall 5985 plantspeciesin a set with
near-minimum
total humanfootprint.The figureshows Africa south of 17QNlatitudewith grey backgroundlines
indicatingnationalboundaries.
530
Annals of the
Missouri Botanical Garden
footprintper cell so thatroadsor populatedplaces
close to relativelyintactareasdo not dominatethe
valuesforthese cells. Fromhereon, meanfootprint
[er cell is referredto as "humanfootprint."
mulativerange-sizerarity."The parameterstested
wereconsideredto be the best measureof ally set
of areas to fulfill the origillalaims of the Myers
hotspots.
Il)k,NTIF\-INC HOTSPOTS ON THE BASIS OF OUR
HOTSPOTS
AS CONSERVATION
PRIORITIES?
DATA
The Myershotspotsare promotedas a network
of areas suitableto "protectthe most species per
dollarinvested"(Myerset al., 2000). This resembles the goal of recentlyappliedheuristicselection
algorithmsforseekingnear-minimum-cost
areasets
(Gaston,1994). The lattertype of area sets represents each species at least once, but tendsto minimize the hypothetical"costs"of potentialconservationactionsby choosingcells with,for example,
least humanimpact (Balmfordet al., 2001; Vlilliams et al., 2003), or at least cost (Mooreet al.,
2004). The approachaimsto alleviateconservation
conflictswherethereis scope for this (Lucket al.,
2004). Technically,the algorithmfor near-minimum-costarea set countsthe numberof rarespecies in each cell (aftertakingfloristiccomplementarity with previouslychosen cells into account)
and then divi(les the diversityscore for each cell
by the humallfootprintvalueof the respectivecell.
Such a near-lllitlimum-cost
areaset was calculated
on the basis;ot the same parametersused for the
hotspotsanalsses. Wethen selected the 125 cells
having the highest benefit-to-costratio and contrastedthem with the Myershotspotsand the redefinedhotspols(Table1, Fig. 2A, B).
(i) Range-sizerarityand humanfootprintwere
normalizecl
to a percentageof theirmaximumvalue,
so that they are bothequallyscaled.
(ii) To rel)resenta measurethat best approximates the iXheat"
of the Myershotspots,we then
calculatedthe productof the value of footprintand
range-sizerarity.This index, which combinesbiodiversityand humanimpact,does not fully reproduce the methodsof Myerset al. and is too simplistic to derive any detailed conclusions on
{onservationpriorities.However,it is suitablefor
tel)resentingthe degree of potentialconflict betweenthe conservation
of diversityandpressureon
lan(l us;eresultingfromexisting humanactivities
(Fig. 1 comparemethodsused by Balmfordet al.,
2001; l uck et al., 2004). Usingthe productensures
that both Xange-size rarity all(l human footprint
tilUSt
11ave(omparativelyhigh values to obtaina
high cotubilledindexvalue.
(iii) The 1713 sub-Saharancells were then
rankedon the basis of the square-root
valuesof the
combinedindexfor each cell.
(iv) The top 125 cells with the highest values
wereselected.Vlechoseonly 125 cells so thattheir
totalareaapproximates
to the totalareaof the Myers hotspots.Fromhere on, the 125 cells selected
on the basis of ourdatawill be referredto as "redefinec]hotspots"(Fig.2A).
(M()MPAIUNGTHE PERFORMANCE OF THE TWO
A1i'I'ERNATIVE SETS OF HOTSPOTS
Althoughthe redefinedhotspotscover the 125
cells with the highestproductof range-sizerarity
alld human footprint,this does not necessarily
meanthat they mustinclude morerarespecies in
total, or indeed a largerproportionof the species'
rangesthando the Myershotspots.Tocompareboth
sets of hotspots we applied three tests: (i) How
manyspecies are included?(ii) Howmanyspecies
belongingto the quartileof species with most-restrictedranges (see Gaston,1994) are included?
(iii) MIhichproportion
of the rangesof the sub-Saharanplantspecies is covered?The lattermeasure
is calculatedby summarizing
the values of rangesize rarityfor all cells includedin each set (analogous to the C-valueof Ikier& Barthlott,2001).
Fromhere on this measureis referredto as "cu-
RESULTS
Thereis a su})stantial
spatialoverlap(Figs.1, 2)
betweenthe Myershotspotsand the redefinedhotspots identifie(lon the basis of our data.The top
17 redefinedhotspotcells are includedin the Myers hotspotsas well. A total of 60 out of the 125
cells coveredbYthe Myershotspotsare also identifiedas redefinedhotspots.
The 125 cells in the Myers hotspotsinclude
3841 of the 5985 sub-Saharanplant species representedin our database(Table1). Fifty-twopercent of the species belongingto the quartileof species with most restrictedranges are included.
Cumulativerange-sizerarityfor all cells is very
high, with 39% of the maximumcumulativevalue
for all 1713 sub-Saharan
cells. The averagehuman
footprintamongthe 125 cells is 24.32; similarvalues are measuredin cells coveringcities such as
Kigali,Kisangani,or Bloemfontein.Valueshigher
than 30 are characteristicof metropolitanareas
such as Durban,CapeTown,Dar es Salaam,Abidjan,and Douala.
.
531
Kuper et al.
Africa's Biodiversity Hotspots
Volume 91, Number 4
2004
TalDle1. Comparative
performances
of three area sets of equal size to cover the sulz-Sahalancenters of plant
diversity.The followingsets are compared:the hotspotsas definedlDyMyerset al. (2000),a redefinedset of hotspots
identifiedon the lDasisof distrilDution
datafor 5985 plantspecieseand a near-ninimum-cost
alea set on the lDasisof
the samedata.In orderto comparethe setseall havelDeenrescaledto a one-degIeelDasedgridof 1713 cells coveIing
Africasouthof 17°N latitude.The comparisonis lDasedon four criteria:(i) osFerall
numlDer
of species covered;(ii)
nwallDer
of restrictedrangespecies covered;(iii) cuIaltllative
rallge-sizeIarityfor the includedcells; and (iv) human
footprint(sumand averageforthe includedcells). Percentagesindicatethe proportion
of the Iespectivetotalsaluesfor
sulz-Saharan
Africa.Theplantdatastemfromthe Biogeographic
Information
Systemon AfricanPlantDiveIsity(BISAP)
representing
of the species of the sub-Saharan
Africanflora.Note thatthe aim of hotspotsets is to cover
the mostthIeatenedcentersof plantdiversity(represented
lDyhighvaluesforlDoth
plantdis-ersity
andhumanfootprint),
whereasthe near-minimum-cost
areaset seeks to coverall species in cells witha humanfootprintas low as possilDle.
1064-156iTC
Sulz-Saharan
Africa
Total
One-degreecells
1,713
All species
5,985
Restricted-range
species
1,540
Range-sizerarity(SUnl)
5,985
Humanfootprint(sutn)
33,965
Humanfootprint(asTerage)
19.8
MyersHotspots
Total
125
3e841
802
2,354
3,040
24.3
Howeverthe Myershotspotsexcludesomeof the
cells with very high range-sizerarityand human
footl)rintand converselyinclude othercelle;with
eoml)aratively
low values for range-sizerarityall(l
humanfootprint(Figs. 1 2). In totale65 cells of
the re{lefinecl
hotspotsdo notc oineide withthe Myels hotspots(*ieel)elow).
In an alea of apl)roximately
the salnesize as the
Myelshotspotstthe redefinedhotsl)otsillelucle15¢Sc
mole of all species and 11%mole of the rareslecies occulling in ourdatabasesfol sub-Saharall
Africa (Table1); in totaltthat is about80% of the
species includedin our databases.Moreovel;they
covera 25.5Wohighercumulativerange-sizerality
than the Myershotspots and they do so in areas
thatare characterizedby a 12.5%highertotalhumantootpllnt.
The redefined hotspots overlap with regional
centersof plantdiversitynot coveredby the Myers
hotspots(Fig. 2A, B). In sollthernAfrica these are
mainlycells in Maputaland,
Pondoland,Barberton
Sekhukhuneland,and Soutpansberg.In Katanga,
the ZambeziSourceArea,Kundelunguand Upemba NationalParkare important.In easternAfrica
the ChimanimaniCentre, Mt. Mulanje,a large
block of cells in the AlbertineRift includingthe
Kivu area,the Ruwenzoriand VirungaVolcanoes,
Bwindiforest, Mt. Elgon, Mt. Kenya,as well as
otherareaswith Afromontane
or Afroalpinevegetation mainly in Kenyaare amongthe redefined
hotspots(Fig. 2). The same is truefor cells in the
lowerGuineaforestblock,includingpartsof southeastern Cameroonand the area between Crystal
Cjtc,
7.3
64.2
52.1
39.3
RedefinedHotspots Near-minimum-cost
set
Total
125
4,759
1,011
2e955
3,420
27.4
Cjto
7.3
79.5
65.6
49.4
Total
125
5,196
1e155
2e603
2,215
17.7
Cjtc,
7.3
86.8
75
43.5
Mts. and the coast in Gabon!the Mayombecenter
of plantdiversity.These areashavebeen identified
as centersof plantdiversits})efore(fot example,in
Brenan,1978; Beentjeet al. 1994; Olson & Dinerstein 1998; van W+k& Smith,2001).
The 125 c ells selecte(l fol the neal-minilnum{ost area set for sul-Saharall[)lulltdiversityhavTe
an ovellal)of 58 cells with the reclefinedhotspots
(Fig. 2). The overlapis high })e{auseall regional
c lusters of ledefined hotsl)otsc ontain cells with
one-cellendertlics.Althoughthe algorithmseeks to
avoid hotspotsand insteaclchose alternativecells
with a low humanimpactethis ie;not possi}le for
manycells becausethey are irreplaceable fol representationof their endemicsl)ecies. Even though
the redefilledhotspotshave been delineatedon the
basis of simplisticcriteriaand methods,the irreplaceabilityof a considerablepalt of theirareaimplies that they will play an importantrole in more
complexand moreadequateprioritizingschemesthereare simplyno alternativesin manycases.
However,the top 125 cells of the near-minimumcost area set include more species and more restricted-rangespecies thanthe Myershotspotsand
the redefinedhotspots,even thoughthey includea
set of cells with a much smallertotal and average
humanfootprint.Bearingin mindthe limitedcomparabilityand simplifiedassumptionsof the algorithm,this result indicatesthat, instead of prioritizingall hotspotsas *'conservation
areas?"thereis
potentialto alleviateconservationconflictsin subSaharanAfricaby prioritizingareas with less hu-
532
Annals of the
Missouri Botanical Garden
manimpactfor conservation,such as, for example, taxa, manyof the newly redefinedhotspotswould
a rainforestblock in centralGabon(see Fig. 2).
certainlyhave an equal priorityin comparisonto
the areascurrentlyincludedin the Myershotspots.
The mostobviousdifferencebetweenthe Myers
DISCUSSION
hotspotsand the redefinedhotspotsis theirspatial
A REDEES1NED
SET OF SUB-SAHARANHOTSPOTS
resolution,in particularfor the Afrotropical
region.
The Myershotspotscover64% of all plantspe- The distributionof the species within the Myers
cies andmorethanhalfof the restricted-range
plant hotspotswas not known,but in our study it was
species includedin our databasefor sub-Saharan possible to optimizethe performanceof the redeAfrica.
fined hotspotson a spatial scale of one degree.
Althoughthis is an impressiveperformancefor Hence, severalcells withinthe hotspotshavebeen
only 125 cells (7.3%of the total area),the rede- replacedby otherspreviouslynot considered.
datalimits
fined hotspotshave an even higherrepresentation The low resolutionof inventory-based
of the threatenedsub-Saharan
flora(Table1). This not only the spatialprecisionof the identification
is mainlyclueto the fact thatsome importantcen- of centersof diversity,but also theircomparability.
tersof plantdiversityunderhighhumanimpactare The Myershotspotsby definitiollhave to illelude
not includedin the Myershotspots.
at least 1500 endemic species per hotspot.This
Severalof these areas have been mentionedby does not take intoaccountthe fact thatsomeareas
Myerset al. (2000),buttheirinclusionin the Myers with high regional concentrationsof restrictedhotspotswasuncertaindue to poordataavailability. range species, such as the AlbertineRift, cannot
These dataare nowavailableand stronglysupport match this thresholdbecause they are too small
inclusion(see also Brookset al., 2002). Moreover, (Plumptreet al., 2003). Since the distributionof
someof the sites havebeen identifiedas majorgaps the endemics within the Myershotspotswas not
in the networkof protectedareas(IUCNcategories knownin manycases, priorityof hots,notswas deI-VI ,lus folest reserves)for threatenedand re- terminedby rankingthe averagenumberof endemstricle(l-Xallge
Afrotropical
plants(Burgesset al., in ics per standardarea.In contrast,taxon-baseddisl)ress). T his emphasizesthe urgencyto consider tributiondata can be used to compareany of the
themill large-scaleconservationassessments.
1713 sub-Saharanone-degreecells enablingcomIn the Myershotspotsanalysiszoologicaldiver- parisonof areassuch as the centralpartof the A1sity served as "backupsupport"(Myers et al., bertineRift or the Maputaland
Centerof planten2000). Patternsof plantandzoologicaldiversityare demismwithparts(notthe average)of the hotspots
not necessarilycongruent.Thereare, for example, in VlestAfrica.In addition,insteadof onlyconsidobviousdifferencesin the importanceof the Vlest- ering the numberof endemicsand the threatper
ern Capeor Kaokoveldin the diversityof restrict- area,taxon-baseddatacan providea varietyof aded-rangeplants comparedto birds or mammals. ditionalparameters,
such as diversityat highertaxNonetheless,the majorityof cells now includedin onomic level or even phylogeneticdiversity,enthe redefinedhotspotsare centersof species rich- ablingcreationof a hierarchyof priorities.
ness andendemismforanimalsas well. Thisis true
for the AlbertineRift (Plumptreet al., 2003) in- TOWAIlD A NETWORK OF PRIORITY ARI"AS FOR
cludingthe Kivuarea,the southwestern
connection CONSERVATION
of the EasternArc to the MbeyaRange,nearlyall
The Myershotspotshad a veryimportant
impact
of the Afromontane
areasmentionedabove,including Mt. Elgon, Mt. Kenya,and the Chimanimani as a pioneeringstudydemarcatingareaswhereonMts., some of the areas in northeasternSouthern going "mass extinctions"(Myers,1990) make it
Africaand also Katanga(Burgesset al., 2004; Cot- most urgentto conservebiodiversity.In parallel,
terill,in press).Range-sizeraritypatternsformam- theyhavebeen promotedalso as a networkof areas
mals, snakes,and amphibians(Brookset al., 2001) suitableto "protectthe mostspecies per dollarinare verysimilar.In analysesthatarecurrentlymost vested" (Myerset al., 2000). At first glance this
since the mostproblemcomprehensive
forsub-Saharan
zoologicaldiversity, seems to be contradictory
manyof these areas have been classifiedas irre- atic areasfor conservationare unlikelyto be those
is mostcost-effective.However,
placeablein the contextof biodiversityconserva- whereconservation
tion (Balmfordet al., 2001) but are inadequately our resultsindicatethat thereare indeedoftenno
protected(De Klerk et al., 2004; Fjeldsa et al., alternativesto protectionof narrowlyendemicspe2004; Burgesset al., in press; Rodrigueset al., cies withincentersof humansettlementand inten2004). If we wantto base hotspotson this list of sive land use. The 125 cells identifiedas redefined
Volume 91, Number 4
2004
Kuper et al.
Africa's Biodiversity Hotspots
533
and hence cannotbe used to generate
hotspotsareinhabitedby morethan80 millionpeo- plementarity
ple (populationdata accordingto [RMIPE],
2002) prioritieson the basis of cost-effectiveness.Tomeet
and the vast majorityof the areas of high plant targetsof initiativessuch as the GSPC,taxon-based
endemismin sub-SaharanAfricaare characterized data are the key information.This includes inforby a very high humanfootprint(Fig. 1). Despite mationon species distributionsin areas that have
their vicinity to metropolitanareas such as Cape so far remainedcomparativelyuntransformed
beTown,Abidjan,Douala,and Dar es Salaam,and cause of difficultaccess and that tend to be undespitebeing oftencompletelysurrounded
by con- dercollectedforthe samereason.Forexample?how
vertedland, manysites of"remainingprimaryveg- many species in the heavily transformedhotspot
etation"(Myerset al., 2000) withinhotspots(such aroundMonroviacould be efficientlyprotectedin
as, for example,Tai and Banco NationalPark of the largely underexploredKrahnBassa National
C6teD'Ivoire,TableMountainNationalParkof the Forestor Sapo NationalPark in southeasternLiRepublicof SouthAfrica,and the CoastalForests beria?Similarly,we do not yet knowthe potential
of easternAfrica)areirreplaceabledue to the many of the undercollectedbordertriangleof Cameroon,
restrictedrange species they contain.Due to this the Republicof Congo,andthe CentralAfricanReirreplaceability,
a largeproportion
of the areasde- publicforthe conservationof the lowerGuinearain
lineatedin the Myershotspotscannotbe substitut- forestand its ecotones.
ed for otherareas, even if the latterhad an even
higherplantspecies richness.In our study,the re- PRIORITIES FOR A GLOBAI
defined hotspots were constrainedby being the CONSERVATION STRATEGY
same spatial size as the Myershotspots.Consegle see threeprioritiesfor the futureapplication
quently,the inclusion of new areas formerlynot
of
taxon-baseddata in the contextof biodiversity
consideredin Myershotspotsresultedin omitting
conservation.
The first prioritymust be to acquire
others previouslyincluded. Moreover,a range of
additionalareascouldnotbe includeddespitehav- these dataat a finerspatialresolution.If the current
ing consi(lerablenumbersof restricteel-range
spe- progressof data acquisitioncolltinues,it is likely
cies and clespitebeing highly threatened.For ef- that before 2010 reliable clistributiondata for
resofective conservation, inclusion of all areas plants will be availableat a quarter-degree
containingclustersof strictendemicsis a millimum lutiollfor the majorityof the globallymostbio{livrequirement.For example,the Namil)clesertand erse ecoregions,a s;calewhich approachesthat at
ari(lwoocllancls
of northeastern
Somaliaarealso key whichactualconservatiollaetionsareimplementecl.
areasfor plant conservation.Moreover,in olelerto In particular,in additionto furtherexplorationof
translatethe hotspotsconceptto conservationin the the hotspots,we urgentlyneecl better information
field,floristicchecklistsof areaswithcomparatively on the biodiversityof remoteareaswithintactvegintact vegetationand, in particular,checklists of etationand of existing l)rotectedareas. A second
existing protectedareas are needed in order to priorityis to combinebotanicaland zoologicalinquantifyhow many restricted-range
taxa are al- formationto obtaina clear pictureof overallbioreadyprotectedso that conservationgaps can be diversity.There is some preliminaryevidence of
determined.This will help Africancountriescom- concordantcentersof endemismamongmanydifply withinternational
agreementssuchas the Glob- ferenttaxaparticularlyin geodiverseareassuch as
al Strategyfor Plant Conservation(GSPC),which Afrotropicalmountains,and these areas should
requiressignatoriesof the Convention
on Biological have a high priorityfor conservationaction.Third,
Diversityto assure protectionof 50% of the most the high resolutionbiologicaldataneed to be comimportantplant areasby the year2010 (Lovett,in paredwith socioeconomicinformationand remote
sensingdataon habitatstatus.Inevitably,biodiverpress).
Althoughmanyirreplaceablepartsof sub-Sahar- sity in largepartsof the redefinedhotspotshas alan hotspotsare in sites of high pressurefor alter- readybeen drasticallyreduced.The combinationof
native land uses, there are also areas with high remotesensing and taxon-basedbiodiversitydata,
range-sizerarityand a comparativelylow human therefore,seems to be a promisingavenueleading
impact(Fig. 1). Theseareascouldcontributeto op- to identificationof those sites wherewe can protect
timizing conservationefficiency and alleviating biodiversityin an efficientand sustainableway.
conservationconflicts.However,the dataon which
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