Patterns of Distribution, Diversity and Endemism of Larger African Mammals
S. Afr.]. Zool. 1994,29(1) 19 Patterns of distribution, diversity and endemism of larger African mammals
J.K. Turpie' and T.M. Crowe FitzPatrick Institute, Universrtyof Cape Town, Rondebosch, 7700 Republic of South Africa
Received 17 November 1992; accepted J June J993
Pallerns of distribution and diversity (= species richness) of larger African mammals, and three subsets thereof (ungulates, carnivores and primates), are identrtied and analysed quantitatively. Distributional patterns generally correspond well with those of presenf.
Diversrteits- en verspreidingspatrone van groot Afrika soogdiere, en die van drie groepe (hoefdiere, vie is vreters en primate), word g'-'dentifiseer en kwantitatief geanaliseer. Alhoewel die verspreidingspatrone goed qo_~~~nst~I'11_r:neU~.Le _v_an_l1u idige plantsoorte~9n>-plaasl ike_nie-akwatiese- yoels; verskil-verspreidi ngsgrense en die graad van verskeidenheid tussen groeps, wat verskillende ekologiese behoeftes weerspies!. Terwyl die
verskeidenheid van groet soogdiere nader aan die ewenaar vermeerder I verskil die patrone van vefskeiden heid tot 'n groat mate tusssn groepe. Daar is 'n positiewe korrelasie tussen soogdierdiversiteit en plantegroei diversiteit, en plantegroei-soorte wat as afsonderlike eenhede g9-analiseer is, vorklaar meer as 85% van die variasie in die soogdiergroepe wat ondersoek is. Verder blyk dit asof beide verspreidings- en verskeiden heidspatrone beinvioed was deur omgewingsfaktore wat in die verlede geheers het.
'" To whom correspondence should be addressed
Any study of biotic distribution, diversity and endemism which occur outside of southern Africa. A 160-quadrat grid requires the identification of pallem before an understanding (Figure I), was used to extract the distributional information of the underlying causal processes can be achieved (Nelson for each species for each quadrat, using a scoring system of . )
9 & Platnick 1981). In this study, we attempt to identify and 0--10 (0 = absen~ 10 = occurring throughout quadrat). The 0
0 interpret patterns of distribution, diversity and endemism for quadrat size was chosen on the basis of software limitations 2 larger African mammals, and focus also on three main
d and was smaller in area than all but 4% of the species' e t subsets thereof: ungulates, carnivores and primates. These ranges. Four mammal data sets were prepared for analysis: a d patterns are then interpreted in the light of past and present all species studied; ungulates (89 species); camivores (60 (
r distribution and diversity of vegetation. The use of vegeta species) and primates (45 species). . e
h tion as the primary factor which influences distribution/
s Vegetational information was extracted from White's i l diversity patterns can be justified as being ' ... the most (1983) vegetation map of Africa. His 80 vegetation types b u meaningful ecological summary of the influences of soil, (including sub-categories) were grouped into 49 broader P climate, topography and other static and dynamic environ e categories, though still adhering to the 17 major groups h t mental factors' (Davis 1962).
described by the author (Appendix 2). For each quadrat. the y
b percentage cover of each vegetation type and the number of
d Methods vegetation types present were recorded. e t
n Raw data a r Numerical and statistical methods
g Distributional information for larger mammals which occur
e in southern Africa was extracted from Smithers (1983). Patterns of distribution and diversity were determined for c n
e Supplementary information for the remainder of the each of the four data sets (all large mammals, ungulates, c i l cominent was obtained from Dorst & Dandelot (1970) and carnivores and primates) using multivariate quantitative r
e Haltenorth & Diller (1977). This study is based on ideal analyses, following the approach employed by Crowe & d Crowe (1982) and discussed in detail by Field, Clarke & n broad distributions of mammals in Africa, i.e. not taking u
into account changes brought about by human settlement Warwick (1982). Cluster analysis, non-metric multidimen y a and associated vegetation changes. Major contractions in sional scaling and information statistic lests were performed w e faunal ranges, however, such as the disappearance of many on the quadrat distributional data in order to identify major t a large mammal species from the south-western Cape (Skead mammalian zones. The Bray-Curtis measure of similarity G
t 1980) are reflected in the distribution maps analysed (Bray & Curtis 1957) and the unweighted pair-group e
n (Smithers 1983). method (Sneath & Sokal 1973) were used in cluster analyses i b In all, 211 species were studied (Appendix I). Insecti to identify any hierarchic similarity among groups of quad a S
vores, rodents and hats were not included in this study, rats. This approach excludes negative matches as evidence y
b owing to the lack of distributional information for species of similarity and takes variation in abundance into accounl, d e c u d o r p e R 20 S.-Afr. Tydskr. Dierk. 1994,29(1)
, between mammal zones. /or-; • ~ A correlation analysis was used to determine the relation ,./. , • • 11 ;{ .. ,J.. between species diversity for the four data sets (number of " " species per quadrat) and the number of vegetation types ~ .. a a\ L" '" " " " " " " present. Stepwise multiple linear regression (Allen 1973) » ~ » M ~ ~ " " " " .. .. .,\ was used to relate species diversity to a combination of ~ ., w ...... ,,'\ vegetation types weighted by their percentage cover of each Q " " ...... a I\r, .. .. • .. -'\ quadrat. Residual plots (deviations from the value predicted " " ",. " ~ " n " n " n " " " ...... 7 by the regression) were used to recognize areas which devi L.!!, ate substantially from the general trends. In these analyses, .. .. ~ ...... ~ ,00) ". " " " those quadrats which had unexpectedly high species diver ,~ ,~ ,~ ,.. 7 " ". '" '" ". sity were considered as possible palaeoecological 'refugia' '" '" '", '" ". '" II( 117/ (sensu Haffer 1969) and, where endemism was also high, ,,. \", '" '" ,1 on ". these refugia were considered to be centres of speciation ,a (i.e. Type" refugia, sensu Crowe & Crowe 1982). ~25 '" ... ". »t n,L 133 ,~ ,,,\ i'" '" '" '" Results ,~ ,., ,~ 'w \'" ,~' V The results for the cluster analyses and MDS for' al1 , ... '.1 • , '\" ,.. .. mammals studied, and each subset thereof, are summarized , p ~ 1~ '" .. ". in Figures 2 and 3 respectively, and the zones of mammal l!e[7 distribution supponed by cluster analysis, MDS and I-tests 'Fe::'"' ./ are depicted in Figure 4. These zones are listed in Tables 1-4, together with information on species diversity, Figure 1 The grid quadrat system used to extract data from the endemism and characteristic species. Patterns of African mammal distribution maps. Superimposed on the grid are the mammal diversity are depicted in Figure 5. Table 5 lists the positions of the major lakes and pans in Africa in the following correlation cocfficient~ between mammal diversity and the grid squares: 63. Lake Chad; 69. Lake Tana; 98. Lake Rudolf, number of vegetation types, and results of the multiple 106. Lake Albert; 112. Lake Leopold II; 116. Lake Victoria; 122. regression analyses between mammal diversity and the Lake Tan7"nia; 130. Lake Malawi; 139. Etosha Pan; 142. Lake different vegetation types are shown in Table 6, The residual Kariba; 148. Makgadikgadi Pan. plots from the correlation between mammal diversity and
. vegetation diversity are sh?wn in Figures 6 and 7. ) 9
0 as opposed to the alternative Jaccard coefficient which is 0 DIscussion 2
sensitive to both presence or absence of species and does d Patterns of distribution e not consider variation in relative abundance (Field & Mac t a Farlane 1968). The validity of quadrat grouping recognized The subregional boundaries of large mammals correspond d
( wel1 with boundaries between major African vegetation by the cluster analyses was assessed by producing a two r e dimensional ordination for each of the four data sets, using types (i.e., forest, savanna and arid zones; Figure 4a). The h s provincial division of the Saharan Subregion represents the i non-metric multidimensional scaling (MDS). MDS is a less l b constraining approach than cluster analysis, since a nonhernmost Iimil for some species (e.g. Addax nasomacu u
P lalis, Crocula crocUla). However, few species are endemic
hierarchic pattern of quadrat similarity is not forced onto the e to this subregion and none are endemic to its Nonhern h data (Crowe & Crowe 1982; sec Shepard 1980). Once the t Province (Table I). The division of the Forest Subregion is y zones (groups of quadrats) were identified, the Information b Statistic test (I-test; Field 1969) was used a posleriori to in{luenced by carnivore and primate distributions, and its d
e Western Province has the highest relative endemism of t determine the characteristic species of each zone. The n larger African mammals. The Savanna Subregion contains
a recognition of distributional zones was dependent on the r 60% of African mammals (Table I) and its subdivision is g quadrat groups being distinct in both cluster analysis and
e in{luenced primarily by ungulate distributions. There is c MDS, and characterized by the presence of several species. n marked variation in the degree of biogeographical zonation e The distributional boundaries of the inost characteristic c i (though not in the placement of boundaries) between l species of each zone (those restricted to and widespread
r ungulates, carnivores and primates (Figure 4), e within the zone) were then used to express the results of the d The extensive radiation of ungulates in Africa is reflected n cluster analyses and MDS canographically. In exceptional u cases, zones were allowed to be delimited by default, i.e, by in the highly fragmented zonation found for this group y a the boundaries of adjacent zones. (Figure 4b). Most ungulate species are found in the Nonhern w
e . Zonal endemics were recognized as those species having Savanna, Southern Savanna and Somali Arid Subregions, of t a more than 85% of their range confined to a particular zone. which the latter has the highest level of absolute as well as G
t The percentage of endemic species out of the total number relative endemism (Table 2). Savannas include a variety of e
n of .mammals in each subset was calculated, and relative vegetation types including bushland, thickets, woodlands i b endemism was calculated as percentage endemism as a and vegetational mosaics, and the Somali Arid Subregion is a
S function of zone area (number of quadrats within that zone), particularly diverse in topography and vegetation, ranging y
b and provides a useful means of comparison, within and from semi-arid lowlands to montane forests and woodlands. d e c u d o r p e R S. Afr. J. ZooJ. 1994.29(1) 21
(a) 100 eo ~ -" 60 "''E '---- '--r- .. '---r- ... 40 Illi II I 20 I 0 (b) II eo €" 0 60 ~ .. -~ ... 40 20 0 (e) (d) III 100 eo eo . ) 9 =E 60 ~ 60 0 -" 0 .~ 2 t d '"... 40 11<---._-, .. '0 e t a d 20 20 ( r e h a a s i l b Figure 2 Large mamma] (a), ungulate (b). carnivore (e) and primate (d) distribution 7.ones suggested by cluster analysis. Roman numerals u P denote subregions and circled numbers represent subregion provinces. e h t y b Only a few smaller species are forest·adapted. e.g. the Genetta genetta) or through most of the area (e.g. Felis d e cephalophines (Bigalke 1972). The incorporation of forest libyca. Herpestes ichneumon). This is partly due to the fact t n savanna mosaics into the Lowland Forest Subregion (Figure that many of the species occur (e.g. Felis libyca) or occurred a r 4b. zone V1.2) is due both to its lack of penetration by many in Europe and/or Asia. The penetration of carnivores into g e savanna species and to its use by lowland foresI species (e.g. desenic areas. the comparatively lower distinction of their c n Cepha/ophus sitvicu/tor). and implies a greater sensitivity to Southwest Arid zone and the absence of a Somali Arid zone e c i all imply a greater ability than ungulates to exploit arid l the presence of forest than for other groups. No species. r areas. and this is probably facilitated by their diet and e however. is confined to this zone. d Many carnivores have widespread distributions refleeting physiology (Bigalke 1978). n u generalized habitat requirements and. consequently. this Primates are largely resuicted in range to the Lowland y a group has fewer distribution zones than ungulates (Figure Forest Subregion. and none occur in the Saharan Subregion w e 4c). With the exception of the fennee (Fennecus zenia) and (Figure 4d. Table 4). In the Lowland Forest Subregion pri t a sand cat (Felis margarita) all carnivores occur in the mates are either distribulCd throughout the zone (e.g. Cerco G t Savanna Subregion (Table 3). and about half are found in pithecus mona. Pan trog/ollYtes). confined to one of the e n the remaining areas. with rclatively fcw species endemic to three provinces [e.g. Cercopithecus diana (\); Cercopithe i b any particular zone. There is some extension of the ranges cus cephus (2); Cercocebus aterrimus (3)J. or in two a S of certain carnivores along the Mediterranean borders of the adjacent provinces [e.g. Cercocebus torquatus (1.2);. y b Sahara. either up the western side (e.g. Mellivora capensis. Cercocebus a/bigena (2,3)J. The remaining species, mainly d e c u d o r p e R 22 S.-Afr. Tydskr. Dierk. 1994,29(1) (a) 69 115-7 140-2 \ 82-85 122-4 145 \,0,' 97-100 129-30 149 1.32-7 ISO /...... 107-109 o .'/ 71-8~""''''''''''' ./ 92-96 CD ..... , 106 1lI 125-7 II n 29-39 VI ~ (b) irA3- 54 !D ( c) II (d) . ) 9 0 n 0 2 m d e CD \0 I t 103-5 a 42 55-85 d 110-4 \" 1~11~2 ( 92-100 118-20 r e h s i l b 151-4 u P ~I e h t ~ y b Figure 3 Large mammal (a), ungulate (b), carnivore (c) and primate (d) distribution zones suggested by multidimensiona1 scaling. Zonal d e symbols as in Figure 2. t n a r g baboons, have extensive distributions, but only three species savannas. There are, however. some notable differences in e c zonation between large mammals and other groups. A n (Papio ursinus. Cercopithecus aethiops and Galago senegal· e c ensi,,) have penetrated the Southwest Arid area. Southwestern Cape Fynbos zone is recognized as a centre of i l r As with the zonation patterns of African birds (Chapin endemism for birds. amphibians and mammals (Bibby et al. e d 1932; Moreau 1952; Crowe & Crowe 1982) and small 1992; Groombridge 1992). but is not defined by large n u mammals (Davis 1962), the major zonational boundaries of mammal distributions. and no large mammals are confined y larger African mammals closely follow those of the major to this area. Similarly. a Montane zone is recognized as a a w vegetation types. The more intricate zonation of Africa by centre of endemism for birds (Dowsett 1980, 1986; Bibby et e t a ungulates corresponds most closely to that for African non· at. 1992), but with the exception of two monkeys (Cercopi· G aquatic birds (Crowe & Crowe 1982). We believe that this thecus l' hoest; and C. hamlyni) restricted to the montane t e relatively fine partitioning of Africa is due to the fact that forests of central Africa. no large mammals are confined to n i b both groups have specialist' which evolved allopatrically in a montane zone. The Dahomey Gap, a break in the lowland a S the same ecological 'islands' of isolated biotopes during forest belt, is reputed to have been an important zoogeogra· y periods of marked expansion and contraction of forest and phical barrier for forest birds (Moreau 1966). However, it is b d e c u d o r p e R S. Afr. J. Zool. 1994.29(1). 23 CD --....•..•... 1--··...•... CD II (a) (b) ." '" II II III . ) 9 0 0 2 (c) (d) d e t a d (b). ( Figure 4 Large mammal' (a), ungulate carnivore (c) and primate (d) subregional (roman numerals) and provincial (circled numbers) r boundaries suggested by the distributions of the most characteristic species. e h s i l b u not significant for larger mammals in general, although it development of a lake, where present mesic forest vegela· P limits the range of certain western forest primales. Within tion exists, served to separate areas to the east and west, e h which during dry phases, became isolated palChes. Crowe & t the Lowland Forest zone, the Cameroon mounlain and high· y lands system acts as a boundary for all the mammalian Crowe (1982) proposed that these areas provided refugia b & (centres of speciation) for forest-dwelling birds. That this d groups analysed, as well as in avifaunal studies (Crowe e t Crowe 1982). This feature is evidently an effective barrier may have also resulted in primate speciation is evident from n a within the forest region, and probably played a role in allo the boundary dividing the central forest region. r g patric speciation. Finally, a grealer proportion of African Also related to past dry periods is Balinsky's (1962) e c large mammals (14/211) than Afrolropical birds (33/1481; proposed arid 'corridor' which linked the arid areas of the n e north-east and south-west. Free migration of animals along c Moreau 1966) have ranges extending beyond the traditional i l nonhern boundary of the Afrotropical Region, into Mediter this corridor would have occurred during relatively arid r e ranean Africa. phases, but during the wetter periods it is postulated that the d n It has been suggested that zonal. boundaries which cannot eastward extension. of the rainforest across the corridor u y be explained by existing physical or environmenlal barriers separalCd animals to the north-east and south-west. This a are related to past environmenlal conditions, and resulting would account for the many disjunct distributions of mam w e t speciation events (Balinsky 1962; Udvardyl969; Bigalke mals (e.g. Madoqua kirkii, Oryx gazelle, Genet/a genet/a) a 1972; Crowe & Crowe 1982; Brain 1985). Historical influ which have populations in the two arid zones, separaled by G t ence on present day distribution of mammals is .evident. areas of moister savanna, as well as the fairly high ende· e n mism found at these extremities. There are also congeneric i Crowe (1978) illustraled hypothetical vegelation patterns b that could have existed during past wet and dry climatic species which are separated in this way (e.g. Equus zebra a S phases. A notewoithy change occurred in the Congo River and E. grevy/), supporting the fact that this process has y b basin within the present Lowland Forest zone, in which the significance as a means of speciation. Moreover, the exist- d e c u d o r p e R 24 S.·Afr. Tydskr. Dierk. 1994.29(1). Table 1 Large mammal zonal diversity, (D), endemism Table 2 Ungulate zonal diversity (D), endemism (E), (E), percentage endemism (%E), relative endemism percentage endemism (%E), relalive endemism (R), (R), and characteristic species. Zone numbers are and characteristic species. Zone numbers are those in those in Figure 4a Figure 4b Characteristic Characteristic Zone name D E %E R marruna]s Zone name D E %E R ungulates I Saharan Subregion 24 ItS 0,09 Gazella dorcas Saharan Subregion 2 0,04 Gazella leptoceros Gazella leptoceros atlZella dMcGS Poecilictus /ibyca II Sudanese Arid 10 4 4 0,16 Dryz donrm4h L Nonhem Province 14 0 0 0 Subregion Gazella dMcGS 2. Southern Province 24 6 3 0,12 Oryx dammilh Addax NUomacuiatis Gazella dama Felis margarita III Somali Arid 35 16 18 1,38 Dryz"""" Subregion RhY'lCotragus glU:nlheri II Lowland Forest 84 41 19 1.12 Perodiclus POliO Lilocran;us wtJJlerl Subregion Galagoitk.t demidovi I. Arid Province 17 4 Cercopithecus mon.a 4 1,33 Gazella 3pe~i AmmodorcGS clarui 1. Westem Province 46 16 8 1.14 Gen.etta pardina Dorcolragws ntl!gaiotis MUllIns gambu:lftUS 2. Highland Province 15 3 3 0,75 Capra ibex Colobus polylUJmos Tragelaplws blalo"; 2. Eastern Province 65 J5 7 0,70 Cepha/ophus flisri/ons EquKr asill.u.s Cephalophw caUipygus 3. Grass Steppe 24 3 3 0,50 Gazella granti Cercocebw albigena Province Damal iscws Jumteri III Savanna Subregion 125 54 26 0,35 Crocwla crocuta TragelapJws ·imberbis Pafll~ra leo IV Western Forest 17 6 7 1,75 CephalopJtus niger Feli.f ,'ifrval Subregion Neotragus pygmtUJIS 1. SomaJi Arid 70 15 7 0,58 Oryx !>eisa CephalopJtws ubra Province LitOCTOllius walleri Gazella soemmeringi v Northern Savanna 28 6 7 0.20 Kobus kob 2. Nonhem Savanna 58 8 4 0,13 Papio an.ubis Subregion CephalopJtus rufdatu.s Province Gozeilo rufifrOllS TalUotragus dermaruu Cepha/ophw ,uJi/alllS . ) VI Central Forest 31 6 7 0,01 CephalopJtus ,"""licola 9 3. Southern Savanna 95 16 8 0.26 lIe/ogole parllula 0 Subregion CephalopJtus SJ/~icu.llor· Province Hippotrogus niger 0 2 CephalopJtus lIigrifrollS Aepycuat melampw d t. Northern Province 21 4 4 0,40 CephalopJtus ClJllypigws e t IV Southwest Arid 52 16 8 0,53 Cyniclws penicillala CephalopJulS lellCogGSle, a d Subregion Vulpe.t chamtJ Cephalophus dorsalis ( r Antidorca.t n'ldr.twpiizlis 2. Southern Province 23 0 0 0.00 e h s i VII Southern Savanna 40 12 13 0,57 Equus blUchelli l b Subregion TalUOiragws oryx u ing forest vegelation associated with lhe montane belt fonns P Aepycerru meJampJlS e an equatorial division of the savanna for all groups except 1. East African h t the carnivores. Province 37 0 o o y b 2. Temperate 32 4 4 0,25 Raphicerus sharpei Fewer camivores have disjunct distributions, however, d Province HippotragJlS niger e and these are separaled by a narrower gap (e.g. Otocyon t Alcelaplws licllleMteill.i n mega/otis) and no sister species arc separaled in this way. a r g This is probably because arid/mesic gradients are not as VII Southwest Arid 25 8 9 0,60 AnlidorCQS marswpiizlis e Subregion Oryx gaulla c effective as barriers for camivores. n Pelet:l capreoillS e c i Patterns of diversity I. Kalahari Province 12 2 2 0.29 Oryx gaulla l r AlcelaphMs caama e Larger African mammals increase in diversity towards lhe d 2. Karoo-grassland 17 4 4 0,50 Pelea capreolws n equator (Figure 5a). This trend has been found for venebl1lte Province Damaliscws @rca u faunas' in Norlh America (Klopfer & MacArlhur 1960; y Raphicerws ntl!lcVWtis a Wilson 1974), lhe fonner Soviet Union (Terent'ev 1963), w e Australia (Schall & Pianka 1978) and for non-aquatic t a African birds (Crowe & Crowe 1982). Even coastal marine G ses have been proposed (summarized in Pianka 1966). t invertebrates and algae follow lhis pattern (Fischer 1960), e The increase in diversity towards lhe equator for North n with ihe exception of burrowing faunas (Thorson 1957) i b which are sheltered from climatic effects. Several workers American mammals is mainly due to the steep increase in a S have atlempled to explain this global phenomenon in lerms bats (Wilson 1974). Wilhout lhis group, quadrupedal mam y b of a universal causal factor, and at least six major hypolhe- 'mal diversity decreases nonhwards only slightly until d e c u d o r p e R S. Afr. J. Zool. 1994,29(1) 25 Table 3 Camivore zonal diversity (D), endemism (E), Table 5 Correlations between percentage endemism (%E), relative endemism (R), . mammal diversity and num and characteristic species. Zone numbers are those in ber of vegetation types Figure 4c Mammal OJaraclerislic groups , p Zone name o E %E R carnivores ALL 0,50 < 0,001 Saharan Subregion 15 0 o 0 UNG 0.49 < 0,001 CAR 0,52 < O,OOt II Lowland Forest 24 13 22 1,29 Felis ilurata PRt 0.20 < 0.011 Subregion Crossarchus ObsCUTUS PoiaNJ riCMrdsoni 1. Western Province 18 3 5 0,71 Guulla pardiNJ Mungos gamiJiaflus group is historically, and thus generally, most adapted. Genella villiers; Ungulate species diversity is greatest in the savanna 2. Eastern Province 19 7 12 1,20 AOIIYx congica woodland areas of the eastern tropical belt (Figure 5b). The Genetla victorioe mosaic structure of this vegetation group, as well as the fine Genetta serva/iM dietary niche separation amongst ungulates, allows the III Saunna Subregion 57 28 47 0,63 Ictollyx striatus coexistence of several species in an area (Lamprey 1963; GelU!11a Seneaa Murray & Brown 1993), and the region is IDpographically Lycaoft piChU highly diverse (Kingdon 1971), incorporating the Ethiopian 1. Savanna Province 35 4 7 0,12 Callis adustus Highlands and African Rift Valley. In addition, it contains He/ogale pal"tluJa the junction of three zones, the sympatry of component taxa Munsos mlUlgo of which could have been overemphasized by the relatively 2. Southwest Arid 19 6 10 0,67 Vulpes chanta coarse scale of data collection. Primate diversity is highest Province Cynictus pefticiliata in the tropical lowland forest (Figure 5d), requiring no Swicala SlI.ricalta further explanation than the presence of their preferred habitat, together with the refugium hypothesis offered earlier. Carnivore diversity is relatively eve,nly spread Table 4 Primate zonal diversity (D), endemism (E), throughout the continent, with the exception of the Sahara, percentage endemism (%E), relative endemism (R), having peaks in the equatorial savanna and the south (Figure and characteristic species. Zone numbers are those in 5c). The southern peak corresponds to the boundary between . ) two zories and may either be an artifact of data collection or 9 Figure 4d 0 a real area of overlap. Camivores are not species-specific in 0 Characteristic 2 their prey choice, although they may be limited (0 a certain d Zone -name o E %E R primates e prey size range (Smithers 1983). Within a biome, habitat is t a Saharan Subregion 0 0 0 0 thus probably important only inasmuch as it affects prey d ( availability, and global trends in diversity do noi, of course, r II Lowland Forest 35 29 66 3,t4 GaJagoides demidovi e apply ID population numbers. h Subregion CercopitMcus mona s i ID l Pemdicticus potlo Crowe & Crowe (1982) attempted explain patterns of b bird diversity in teons of vegetation diversity (number of u 1. Western Province 22 5 tt t.83 C oIobus po/y/wmos P Cerc.opithecus petaUTista vegetation types). Analyses of this kind in the case of e h Co/obus vents mammals revealed even weaker correlations than found for t y 2. Centra1 Province 26 10 23 4,60 Euoticus e/ega1l1u/us birds (Table 5). However. vegetation types analysed as b GaJago alieni separate units in multiple regression analyses were able ID d e ArcJocebus CAllabare1lSis t explain over 85% of the variance in mammal diversity in all n 3. Eastern Province 28 3 7 0.70 CoJobus allgole1lSis cases (Table 6). The importance of desert as an inhibitory a r Cercopithecus ascallius g factor is emphasized, but the apparent negative effect of e Colobus peManti c Mediterranean selerophyll forest may be due to its inaccessi n e III Savanna Subregion 8 5 11 0,22 GaJago sefll!gaJe1lSis bility. Tropical vegetation types are an important correlate c i l Cercopithecus aelhiops of species diversity, and the major positive influences for r Cercopithecus mitis e each mammal subset are the vegetation types associated d n with their ecological requirements. Thus, it follows that u most variance in primate diversity is associated with the y a extreme conditions north of 50"N are encountered. In least amount of vegetation types, and deserts are only of w e comparison, the increase of larger African mammal diversity secondary importance to carnivores. t a IDwards the equaIDr is steeper, but there are facIDrs specific Much of the deviation in diversity trends (Figures 6 and G t ID each major group which can probably account for much 7) can be explained by the presence of landscape features e n of this gradient such as water bodies and mountains. In all the groups i b The'region of greatest diversity for each mammal group studied, lower than expected diversity is found in the Marra a S mountain area of Sudan and, except for primates, in the in Africa is not distributed across the whole equatorial y b region, but is restricted to the vegetation type ID which each Ethiopian and Somali lowlands. Low diversity associated d e c u d o r p e R ,',. 26 S.·Afr. Tyd,kr. Dierk. 1994. 29(lr', Ca) '0 ~20 . ) 9 0 0 2 d (e) Cd) e t a d Figure 5 Geographical variation in diversity of the larger African mammals (a), ungulates (b), carnivores (c) and primates (d). ( r e identify areas of importance for the conservation of biotic h s i with Lake Victoria is probably due to the corresponding gap diversity. Thus, in the lowland forest, the central province is l b on the distribution maps created by the lake. Carnivores also the centre of highest primate endemism. and the western u P have lower than expected diversity in the Atlas Mountains province is important for all of the three subgroups. The e h of Algeria, the Guinea highlands and Cameroon Highlands. Somali Arid Subregion is the area of highest ungulate t y Higher than expected diversity, in all groups except endemism and carnivore endemism is fairly high in the b primates, is associated with rivers or lakes in arid areas: Southwest Arid Province. The areas richest for endemic d e Lakes Chad. Tana, Tanganyika, and the East African lakes; t mammals (including small mammals) identified by Bibby et n the Nile River, the Niger River and associated wetlands in al. (1992) include these above-mentioned areas as well as a r g central Mali, and the Okavango River. Higher than expected montane and Cape fynbos areas. However, although this e c diversity for all groups except primates is also found in the endemism-based approach has been used by Bibby et al. n e vegetational mosaics south of the lowland forest, in the (1992) and others to identify and prioritize areas for c i l mountainous area around Swaziland, and in Namaqualand. conservation. these areas do not coincide with the areas of r e Primates have lower than expected diversity in the central highest diversity for larger mammals. which occur in the d lowland forest. in the region of the Congo River and savanna regions, except in the case of primates. As an n u associated marshlands and lakes, and higher than expected additional criterion. the degree of zonation within each y a diversity in the lowland forest immediately to the west of group serves as a guide to the variety of areaS that need to w e this area. These patterns further support the idea of a central be conserved in order to preserve maximum diversity, and it t a forest barrier and adjacent refugia. Higher than expected is further necessary to identify areas of high endemism and G primate diversity is also associated with highlands in richness within these particular biogeographical regions t e Guinea, Cameroon, and east Africa. (Rebelo & Siegfried 1992). Thus, the conservation of n i b representative taxa for each subregion is a minimum a S Implications for conservation requirement. and within each biogeographical province, an y b Centres of endemism and diversity are two criteria used to ideal goal. d e c u d o r p e R S, Afr, J, Zoo1. 1994,29(1) 27 Table 6 Summary of results of stepwise multiple regression analyses of mammal diversity vs vegetation diversity (VEG), The ten best correlated vegetation types and their contribution to the total multiple coefficient of determination (R2) are listed for each group AU species Ungulates Carnivores Primates-_ Sign Sign Sign Sign VEG· R' of, VEG R' of, VEG R' of, VEG R' of, 70 0,36 70 0,31 25 0,32 + 0,42 + 67 0,19 67 0,12 10 0,14 II 0,20 + 45 0,05 + 45 0,09 + 67 0,13 25 0,10 + 25 0,04 + 25 0,06 + 19 0,05 + 8 0,06 + 42 0,04 + 42 0,07 + 42 0,04 + 19 0,06 + II 0,03 + 19 O,DJ + 22 0,03 + 42 O,DJ + 19 0,03 + 10 O,DJ 34 0,02 + 15 O,DI + 10 0,02 0,02 + 10 0,02 35 0,01 + 35 0,02 + 35 0,02 + 58 0,01 + 45 0,01 + 0,02 + 38 0,02 + II 0,01 + 40 O,DI + Tol. R' = 0,90 Tot. R2= 0,86 Tol. Rl = 0,86 . To!.. R2 = 0,89 • I = Lowland rain forest; 8 = Swamp forest; 10 = Mediterranean scleryphyllous forest; 11 ='Lowland forestlSecondary grassland mosaic; IS" = West African coastal mo§sic; 19 = Montane vegetation; 22 = Forest I secondary grassland I woodland mosaic; 25 = Woodland; 34 = S.A. scrub woodland I High· veld grassland mosaic: 35 = W Conclusions mammals, but the degree of zonation within each group pro Patterns of distribution vary amongst the main groups of the vides a guideline as to the complexity of reserve networks . larger African mammats, Distributional boundaries tend to required for the conservation of maximum diversity, ) 9 be physical or ecological barriers which, in most cases, 0 0 ACknowledgements 2 closety fottow the timits of major vegetation types, Where d vegetation fonns a gradient of gradual change from one type We are grateful to L.G, Underhill for kindly attowing us to e t a 10 another (e,g, savanna 10 lowland forest), boundary limits, use (and for adapting) his MDS program for the original d ( as dictated by the ecologicat requirements of the group, are honours projcct (Department of Zoology, University of Cape r e 'flexible' to a degree which depends on the steepness of the Town), h s i gradient. l b Although the zones identified correspond to a certain References u P extent to the major biotic zones that have been identified by ALLEN, J.A. 1973 SmPREG 1: Stepwise multilinear regression, e h other biogeographicat studies, it has become evident that to analysis. University of Wisconsin, Madison. t y attempt 10 define these boundaries as a steadfast rule is RALINSKY, B.I. 1962. Patterns of animal' distribution of the b d unrealistic, The heterogeneity of the various faunal compo African continent. Ann. Cape Provo Mus. 2: 229-310. e t nents detennines some major differences, and even within BlBRY, C.J .. COLLAR, N.J" CROSRY, M.J .. HEATH, M,F.. n a the mammals, there are major differences between groups, 1M RODEN, Ch" JOHNSON, T,H .. LONG, AJ .. r g Patterns that cannot be explained solely by existing environ STAlTERSRELD,AJ, & THIRC.ooD, S.J, 1992, PUlling e c biodiver~ity n mental features can mostly be attributed to vegetational on the map: priority areas for global conservation. e c changes that have occurred in the past. The patterns of International Council for Bird Preservation. Cambridge. i l r diversity and endemism observed substantiate much of the BIGALKE, R,C, 1972, The contemporary mammal fauna of e d evidence for these evolutionary events, Africa. In: Evolution. mammals. and southern continents. n u Reasons for the equalOr-ward increase in diversity (Eds) Keast, A" Erk, F,C, & Glass, R, State University of y New York Press, Albany. a become etearer when separate mammal groups are observed, w This trend is largety due to the concentration of primates in RIGALKE, R,C. 1978, Mammals, In: Riogoography and ecology e t a the equalOrial lowtand forest. and to the radiation of of Southern Africa, (Ed,) MJ,A, Werger, pp, 981-1048, Dr G ungulates in the vegetationatty and topographically diverse W. Junk. The Hague . . t e n regions of East Africa, BRAIN, C.K. 1985. Tcmperaturc-enduced environmental chang~ i b Finally, the discrepancies in' areas of high endemism and in Africa as evolutionary stimuli. In: Species and speciati,on a S diversity between and within the different subseL' compli (cd.) E.S. Vrba. pp. 45-52. Transvaal Museum. Pretoria. y b cate the identification of conservation 'hotspoL" for large BRA Y, J,R, & CURTIS, J,T, 1957, An ordination of the upland .. d e c u d o r p e R 28 S.·Afr. Tydskr. Dierk. 1994, 29(1)' ..: (a) (b) -2 , -2 -, , " " " 20 30 30 " " " ==j_======__ ~l " -"'------= " " 90 ., '" 100 -2..~ ______ . ----,-~ ) 9 0 '" 0 '" 2 ~ ______1.!i d ------0 e t '" ______iJ a '" d ( r >3, e h '" s i l b u P '" '" e h t y b '" '" d e _-.J------",l t n UK. ,eo a r g Figure 6 Standardized residuals (Allen 1973) for each sampling quadrat for large mammal (a) and ungulate (b), diversity liS vegetation e c diversity regressions. n e c i l r e forest communities of southern Wisconsin. Ecol. Monogr. 27: Muridae. with notes on some of their fossil antecedents. Ann. d n 325-449. Cape Provo Mus. 2: 56-76. u CHAPIN. J.P. 1932. The birds of the Belgian Congo. Part J. Bull. DORST, J. & DANDELDT, P. 1970. A field guide to the larger y a Am. MusewnNalur.lJisl. 65: 1-756. mammals of Africa. Collins, London. w e CROWE, T.M. 1978. The evolution of guincafow] Galliformes. OOWSErI'. R.I. 1980. Extinctions and colonisations in the t a (Phasianidae, Numidinac): taxonomy, phylogeny, speciation African montane island avifaunas. Proc.lV Pan-Afro orn. G t and biogeography. Ann S. Afr. Mus. 76: 43-136. Congr.185-197. e n CROWE. T.M. & CROWE, A.A. 1982. Patterns of distribution, DOWSErI'. RJ. 1986. Origins of the high-altitude avifauna of· i b diversity and endemism in Afrotropical birds. J. 7..001 .. Lond. tropical Africa. In: High altitude tropical· biogeography. (cds) a S 198: 417-442. F. Vuilleumicr & M. M_onasterio. pp. 557-585. Oxford y b DAVIS, D.H.S. 1962. Diso-ibulion patterns of routher~ Africa University Press, Oxford. d e c u d o r p e R S.Afr.J.7.ooI.1994,29(1) 29 (a) (b) -2 -, • -2 -, • .. .. 3. 3. •• -l==:-'-=.---'"' •• ______.1.~ ------~oi •• •• •• •• -'''----==---~ !.Q_------7. ______:!. ..r! ~------•• •• to -i.~ __ . ______00 ------~:~ "'. ,.. ~J! . ______) 9 0 ______19 0 n. 2 ". ~.g ------d _. ______},!i e t ,,. a d ( r e , h .. 03. s i l b u , P .. e h t y , b .. ". d e t n ,eo a ,eo r g Figure 7 Standardized residuals (AUen 1973) for each sampling quadrat for carnivore (a) and primate (b) diversity ",s vegetation diversity e c n regressions. e c i l r e FIELD. 1.0. 1969. The use of information statistics in the Evol. 14: 64-81. d n numerical classification of heterogenous systems. 1. F:col. 57: GROOMBRIDGE. B. (ed.) 1992. Global bindiversity: Slatus of u 565-569. the earth's jiving resources. Compiled by the World y a HELD. 1.G .. CU.RKE, K.R. & WARWICK, R.M. 1982. A Conservation Monitoring Centre. Chapman & Hall, London. w e practical strategy fOT analysing multispecies distribution HAFFER,1. ]969. Sp:cciation in Amazonian forest birds. Science, t a patterns. Mar. Eeol. Prog. Sey. 8: 37-52. Warh. 165: 131-137. G HELD, J.G. & MACFARLANE, G. 1968. Numerical methods in HALTENORTH, T. & DILLER. H. 1977. A field guide to the t e marine ecology. 1. A quantitative 'similarity' analysis of mammals of Africa. Collins, London. n i b rocky shore samples in False Ray, South Africa. Zoo1. afro 3: KINGDON, J. 1971. East African mammals. Academic Press. a S 119-137. London. y FISCHER, A.G. 1960. Latitudinal variation in organic diversity. b KLOPFER. PH & MACARTHUR. R.H. 1960. Niche size and d e c u d o r p e R 30 S.-Afr. Tydskr. Dierk. 1994,29(1) faunal diversity. Am. Nal. 94: 293-300. Appendix 1 African mammal species used in this study LAMPREY, H.E 1963. Ecological separation of the large PR1MATA CARNlVORA mammal species in the T arangire Game Reserve. Tanganyika. E. Afr Wildl. J. 1: 63-92. Perodicticus poliO Prolelu crislaJus HyaellLJ brWlMa MURRA Y, M.G. & BROWN, D. 1993. Niche separation of Artoc~bus caiabar~nsis Galago alieni HyaellLJ hya~1ILJ grazing ungulates in the Serengeti: an experimcnral test. 1. Galago crassical4datus CrocuJa CTocwta Anim. Eco[. 62: 380-389. Galago senegalensis Acinonyx juoolws MOREAU, R.E. 1952. Africa since the Mesozoic: with particular Ewoticus elegafIJulus Panth~ra pardus PantMra leo reference to certain hiological problems. Proc. 7.001. Soc. Galagoities thmiJoyj F~/is caracal Lond. 121: 869-913. Papio lsamadryar F~/is libyca MOREAU. R.E. 1966. The birdfaunas of Africa and il~ islands. Papio papio Felis nigrjp~s Academic Press. New York. Papio anwbis F~/is serval NELSON, G. & PlATNICK, N.!. 1981. Systematics in Papio ge/ada F ~/is mmgarita biogeography: cladistics bicariance. Columhia Universil}' Papio sphinx F~/is allTata Papio I~wcophaeus Press. New York. Felis chaus PIANKA. E.R. 1966. Latitudinal gradients in species diversity: a Papio wrsinus Otocyon megalOlis review of concepts. Am. Nal. 100: 33--45. Papio cynocephalus Lycaon pictus Cercocebus torqwalus REBELO, A.G. & SIEGFRIED, W.R. 1992. Where should nature Vulpu cMma Cercocebus gal~rjlus reserves be located in the Cape floristic region. South Africa? Vulpes "wIpes Cercocebus aJerrimus Vulpes ruppelli Models for the spatial configuration of a reserve network Cercoc~bus albigena Vulp~s pail ida aimed at maximising the protection of floral diversity. Allenopilhecus nigroyiridis Canis adustus Consav. Bio[. 6: 243-252. Miopithecus lafapoin Canis f7U!Somelas SCHAll, JJ. & PIANKA, E.R. 1978. Geographical trend, io C~rcopithecus cephus Canis aur~us numbers of species. Science 201: 679-685. Cercopithecus erylhrotus Canis stmensis SHEPARD. R.N. 1980. Multidimensional scaling. tree-fitting, Cercopithecus erylhrogast~r Aonyx capensis and clustering. Science 210: 390-398. Cercopithecus dfcanius AOtIyx congica SKEAD, C,J. 1980. Historical mammal incidence in the Cape Cercopithecus p~taWTisla LuJra mac'ulicollis Mel/iyora capeflSis Province. Department of Nature and Environmental Cercopithecus njctitans Cercopithecus milis Poecilogale albin.ucha Conservation. Cape Town. Cercopithecus mona /ctony:r. striiltus SMITHERS. R.N. 1983. The mammals of the southern African Nandiniil bil10tata . Cercopithecus diana ) subregion. University of Pretoria, Pretoria. 9 Cercopithecus Mg/~ctus CiYetlictis civelta 0 SNEATH, P.H.A. & SOKAl, R.R. 1973. Numerical taxonomy. G~nelfa g~Mtta 0 Cercopithecus /' hoesti 2 W. H. Freeman. San Francisco. Cercopithecus Mmlyni Genetta tigrin.a d e TERENT'EV, P.V. 1963. Attempt at application analysis of pyg~rylhrus Genelfa pardillLJ t Cercopithecus a variation to the qualitative richness of the fauna of terrestrial Cercopithecus albogwlari.f Genetta victoritu d ( Genelta atryssinica vertebrates of the U.S.S.R. Veslnik uningrad~k Univ. Ser. Erylhroc~bus paw r e Genetta yil/iersi Colobus yerus h Bio!' 18: 19-26. English abstract in BioI. Abstr. 80822. s G~netla servalin.a i Colobus badius l THORSON. G. 1957. Bottom communities (sublittoral or shallow Suricata sWTicatta b Colobus pennanti u shelO. In: Treatise on marine ecology and paleoecology. (Ed.) Paracyniclus selousi P Colobus polykombs H.S. Ladd. Geoi. Soc. Al1ll'r. Mem. 67: 461-534. CYllictus penicillo.ta e Colobus ango/~nsis h UDVARDY, M.D.F. 1969. Dynamic zoogeography. Van t Herpestes icJiMwnon Colobus atryssinicus y Nostrand Reinhold Company, London. Herpestes naso b Colobus satana.s Galerelia sallguinea d WHITE, F. 1983. The vegetation of Africa. UNESCO, Gorilla gorilla e t Galuella pwIverwIenta Swit7.erland. Pan lroglodyles n Rhynclwgale mellen' a 'NILSON, 1.W. 1974. Analytical zoogeography of North r Ichl1.l!wmla aibica~ g American mammals. Evol. 28: 124-140. PIlOLlDOTA e Atilax pal/ldinosus c n Manis t~mminclci Mwngos mullgo e c Manis (SnwLsiil) gigafIJia MlI.ngos gambiilnus i l Manis (Phaloginus) tricuspis H~/ogal~ parvultJ r e Manis (Vromanis) tetradaClyla F~nnecus urdiJ d n Poecilictus libyca u PoiilllLJ richardsoni y LAGOMORPHA a Pronolagus rupeslris Osbornictws pisciYora w e Bd£odiJle crassicauciaJa t Pronolagws crassjcal4dat/L~ a Pronolagu.f ra1!deflSis Bd£ogale njgripes G t Bunolagus monJicwlaris Dologale dybowskii e Crossarchus obscurus n upus capensis i b upus sax.a.tjlis a S TUBULIDENTATA upus crawshayi y b Poelagus marjorita Orycuropus afer d e c u d o r p e R S. Afr. I. Zoo1. 1994, 29( 1) 31 Appendix 1 Continued Appendix 1 Continued PROBOSCIDEA AluJaphw Jichlenslt.ill.jj Raphicerw melaflOlis Kobus ledle A/aJapnus busepltalllS Raphic~rus sharpei Kobus lIlJrdon.ji Lozodollla aft icallQ DamaJiscus dorcas N~otragus moschalus Kobus defassa DamaIiscllS funatus N~otragus pygma~us HYRACOIDEA K obus megac~ros Damaliscws kbrrigllm N~otragus btJt~si Hylochoerw: mLineTlzhagelli ProctlviLJ cap~f!l.Sis Damaliscus hunteri A~pycuos m£lampus Ammo/ragw teTVia HtteroJryrax brlilai Cephalophus monlicola Pelea capTl!olw: Okapia jOMs/olli Dur.dTohyrax. (uOOrt.u.5 Cephalophus nalaiensis lIippolragus ~quillw Boocerw euryceros Cephalophus syJ\licultor Hippolragus n.ig~r Addax NJSOf7tlJcuJalUS UNGULATA Cephalophus ~lItilllU Oryz gaz~lIa Lilocrallius walleri Ct.ralolnuirun sinulm Cephtllophus spadix. Oryr dtlmmo.h Afnlnbdorcas clmJcei DiceTos bicorllis Ct!phalophus niger Oryx beisa Gaul/a dnmtJ Cephalophus TUji/tJ/US Syllcerus ca/Jer Equus ub,a Gaul/a soe~rillgi Equus burclu.JJi Cephalophus zeb"l Trag~laphus strepsiaros Gazella grallti EqUIlS (Asinus) osinus Cephalophus callipygus Tragelaphus speui Gaze/Ja dorcas Trag~laphus QIIgasii EqUIlS (DalidwhipplLf) grevyi Cephalophus dorsalis Gazella peizellli Trag~laphus scriptw CJweropsis libu~1ISis CephaJophus leucogasler Gazella speui Tragelaphus imberbis Sus seTa/a CephaJophus oIgibyi Gazella rufifroll.S PhaCOCJt.oe'1Lf atthiopicus Cepha./opln.u "igrijroflS Trage/aphus bU%lOlli (;au/Ja thomsolli PotamocnOeTUS porc/,4,S SY/lliCLlpra g,immia TaWOlragw oryx Gazella leptoceros lIippopOlamlJ.S amphibiu.t Antidorcm marsupuJlis TaWOlragus derbiallus Dorcolragus mega/otis H~mmchw.s aquaricus O,eotragus ort!otragus Redun.ca a"mdinum Rhyn.cotragus gu.enlheri Giraffa camelofXlrdolis Madoqua kirJd.i RedulICQ fulvorufula Maeioquo sallioNl COftMCNUles gnow Our~bia our~bi Redun.ca ndufICa Capra ibex COftMChaetes tauTinus Raphicerus CQmp~.'dris K obw ellipsiprymnus Appendix 2 Grouping of White's vegetation types and description of each group White's Major type map units Description . ) 9 Forest 1·3 Lowland rainforest 0 0 4 Transitional rainforest 2 d 6 Zambc7jan dry evergreen forest e t Forest transitions 11-14 Lowland rainforest and secondary grasslands a d and mosaics 15,16 Coastal mosaics ( r 17 Cultivation and secondary grassland replacing upland and montane forest e h 19 Montane vegetation s i l 20 Transition from Afromon1.8ne scrub forest lO Highvcld grassland b 21 Mosaic of Zambesian dry evergreen forest and wetter miombo woodland u P 22 Mosaic of dry deciduous forest and secondary and wooded grassland e h 23 Mosaic of Mediterranean montane forest and ahimontane shrubland t 24 Mosaic of AfromonLane scrub forest, 7...amhczian scruh woodland and y b secondary' grassland d e t Woodland 25-30 Woodlands n a r Woodland mosaics 31 Mosaic of wetter 7..ambc7~an woodland and secondary grassland g Jos and Mandara Plateau mosaics e and transitions 32,33 c ~crun n 34 Transition from South African woodland to Highveld grassland e c 35 Woodland transition to Acacia deciduou!> bushland and wooded grassland i l 36 Transition from Colophospun'Ul.m mopa1l£ scruh woodland to r e Karoo-Namih shruhland d n u Secondary wooded 37 Acar.ia polyactJn1ha secondary wooded gra!>sland y grassland a w e Bushland and 38,39 Evergreen and semi-evergreen bushland and thickct t a thicket 40 Itigi deciduous thicket G 42 Somalia-Masai deciduous bushland and thicket t e 43,44 Deciduous wooded grassland and bushland n i b Bushland and 45 Mo,o;aic of Ea~l African evergreen hushland and secondary Acacia a S thicket mosaics wooded grassland y b 47 Mosaic of Brachyslegia btJlcualfQ. thicket and cdaphic grassland d e c u d o r p e R 32 S.-Afr. Tydskr. Dierk. 1994.29(1) Appendix 2 Continued White's Major type map units Description Traditional 48 Tugela basin wooded bushland scrubland 49 Transition from Mediterranean ArgGlnia scrubland to succulent semi-desert shrubland Cape shrubland 50 Fynbos Semi-desert 51 Bushy Karoo-Namib shrubland vegetation 52 Succulent Karoo shrubland 53 Dwarf Karoo shruhland 54.55 Semi-desert grassland and shrubland 56 The KalahariIKaroo-Namib transition Grassy shruhland 57 Grassy shrubJand Grassland 58 Highvcld grassland 59";; I Edaphic grassland Edaphic grassland 62 Mosaic with Acacia wooded grassland mosaics 63 Mosaic with communities of Acacia and broad-leaved trees 64 Mosaic with semi-aquatic vegetation Altimontane 65.66 Altimontane vegetation Desert 67";;9.74 Vegelalioniess deserts 70-72 Dcscns with vegetation A7,onal 75 Herbaceous swamp and aquatic vegetation 76 Halophytic vegetation 77 Mangrove 78-80 Anthropic landscapes . ) 9 0 0 2 d e t a d ( r e h s i l b u P e h t y b d e t n a r g e c n e c i l r e d n u y a w e t a G t e n i b a S y b d e c u d o r p e R