B othalia 15. 3 & 4: 613-629 (1985) Analysis of the size and com position o f the southern African flora G. E . G IB B S RUSSELL* keywords; endemic spccies. flora, phy(oleography, plant families, species diversity ABSTRACT The southern African flora has been surveyed for the first time at spccies level in the List o f Species o f Southern A frican Plants (Gibbs Russell e ta l., 1984). The numbers o f taxa recorded for southern Africa are compared to the numbers reported from other parts o f Africa, and the largest families in each area are listed and compared The species richness o f southern Africa is compared to that o f o th er parts o f the world. The numbers of genera, species and infraspecific taxa are given for each familv in the souihern African flora, and compared to previous counts bv D ver (1975. 1976) and G oldblatt (I97H). INTRODUCTION T he rem arkable diversity and high level o f e n d e mism in the southern African flora has been re ported by a n um ber o f authors, notably A dam son (1938). W eim arck (1941), Levyns (1964) and G ood (1974). and has been discussed in great detail by G oldblatt (1978). U ntil recently there was no m od ern inventory o f the taxa on the subcontinent, but the publication o f the List o f Species o f Southern African Plants (G ibbs Russell et al.. 1984) now p ro vides for th e first time a com plete coverage o f the en tire flora. T he com ponents of the southern Afri can flora can now be precisely analysed and com parisons can be m ade with floras o f o th er parts o f A frica and with previous estim ates o f the southern A frican flora. T he relative im portance o f present studies can be assessed, and future w ork o n the Flora o f Southern Africa can be planned w ith a m ore accurate idea o f the m agnitude o f the task, M ETHODS T h e num bers o f taxa reported here for southern A frica are taken from the P R E C IS list o f 30 June 1984. and som e m odifications have been m ade since the first edition o f the List o f Species went to press in O c to h c r 1983. T he taxa in the southern A frican flora w ere co u n ted by com puter, and have been verified by a m anual count o f the List o f Species. C ounts for o th er Floras in Africa were m ade by hand, as d e scribed in G ibbs Russel! (1974), Because the Floras considered for this study differ in th e delim itation o f families and in the level o f re cognition o f species and infraspecific tax a. it was necessary to adopt a uniform treatm ent in o rd e r to com pare them . Families are treated sensu lato , and th e genus and species counts for the segregate fami lies are added to give a single count in these cases. N otable exam ples o f families treated in this way arc A izoaceae (includes M esem bryanthem aceae). Asclepiadaceae (includes Periplocaceae), C am panulaceae (includes Lobeliaceae), Fabaceae (includes C aesalpinioideae. M im osoideae and Papilionoid e a e ), Liliaceac (includes A lliaceae, A sparagaceae, * Botanical Research Institute, Department of Agriculture & W ater Supply. Private Bag X101. Pretoria 00(11. A sp h o d e la c e a e , C o lc h ie a c e a e . D ra c a e n a c e a e , Eriosperm aceae and H yacinthaceae) and Scrophulariaceae (includes Selaginaceae). T he alternative to a broad acceptance o f these families w ould have been to split them apart in Floras which treat them as units. This was not done because an object o f the study is to convey an overall picture o f the southern African flora in relation to the floras of o th er parts o f A frica, and it was therefore m ore consistent to accept these families in the broadest sense. It was also necessary to adopt as far as possible a uniform treatm ent o f the low er tax a, especially those o f infraspecific rank. Because one author's species may be another author’s subspecies o r variety, the total species num ber for different accounts o f the sam e group can differ considerably, and are not readily com parable. For this reason, com bined totals o f species plus infraspecific taxa w ere used o n the tables below' for purposes o f com parison, For ex am ple, Crassula was revised for southern Africa by T olken (1977). and this revision is followed in the List o f Species. There are now 237 taxa, including 142 species with 47 subspecies, 39 varieties and 9 re cognized hybrids. Exam ination o f his treatm ent shows that the great m ajority o f taxa now accepted by him at the infraspecific level w ere originally re cognized as species. O f the 29 species given for the C ape Peninsula by A dam son & S alter (1950). 9 are accepted at the infraspecific level by T olken (1977). If these treatm ents were com pared a t the level o f species, 30% o f the Crassula species in A dam son & Salter (1950) would not be counted in the List o f Species that follows T olken, thus giving the A dam son & Salter (1950) count a falsely high com parative value. Just as the different Floras considered here cannot be directly com pared because o f differences in tre a t m ent, so the different parts o f the List o f Species it self vary widely in their ranking o f taxa because the list is the result o f taxonom ic judgem ent by n u m er ous individuals m ade over at least 80 years. A re cently revised group such as Crassula may contain fewer species and m ore infraspecific taxa. while a group greatly in need of revision, such as the entire family M esem bryanthem aceae, presently has a great m any species that will probably be reduced when they are critically studied. T he num bers o f species 614 B othalia 15, 3 & 4 (1985) and infraspecific taxa is therefore used as a conserva tive ‘low est com m on denom inator' m aking it pos sible to trea t at m ore o r less the sam e level Che floras o f different areas studied at different tim es by differ en t individuals. taxa once a basic list is p rep ared , the total num bers can in fact be calculated in a num ber o f ways d e pending o n the em phasis th at is required. Infraspe cific, naturalized and well-known but as yet unpub lished taxa may be included o r excluded. T he two m ost extrem e o f these different totals are used below to m ake com parisons with the floras o f o th er areas. T he to tal num ber o f species and infraspecific taxa (including naturalized tax a, unpublished taxa, sub species, varieties and recognized hybrids) is a ‘high’ total (Table 1), and the num ber o f indigenous species is a ‘low ’ total (T able 2). T he problem o f outright synonym y has been im possible to solve w hen w orking a t a continental scale o v er a tim e span o f several decades. T he ideal would be to work through each o f the Floras considered w ith th e aid o f th e most recent taxonom ic revisions and thus ensure that each taxon is recognized in the sam e way. H ow ever, this would be equivalent to re vising the entire A frican flora before m aking the com parisons presented here. Because this is totally im practicable a t p resen t, each o f the Floras covered is taken as it stands and the taxa accepted by the au th o r o f each treatm ent are accepted in this study, even though som e o f the taxa counted have been, or should have b een , placed in synonym y. T he use of species and infraspecific taxa w hen m aking the com parisons eases som e o f the discrepancies introduced by different levels o f treatm ent a t different times, but cannot elim inate them, T h e total num ber o f species and infraspecific taxa is used to com pare the Floras o f different areas, and different parts o f the List o f Species, fo r the reasons o f inconsistencies in level o f taxonom ic treatm ent given above. T he ‘high’ total is also used to m ake estim ates o f the outstanding w ork for the Flora o f Southern Africa because all infraspecific and natura lized taxa must b e dealt with. In practical term s both o f these can be tim e-consum ing to cover for the Flora. T he total num ber o f indigenous species m ust be used for biogeographical studies because the natura lized aliens, which have only recently becom e part of the African vegetation, are elim inated. This 'low ' count is used to calculate species/area ratios to com pare the diversity o f the southern A frican flora to that o f o th er parts o f the world. It is also used to RESULTS AND DISCUSSION Total numbers o f taxa in the southern African flora T he num bers o f taxa present in southern Africa are shown in T ables ! & 2, A lthough it may appear to be a sim ple m atter to determ ine total num bers o f TABLE 1, — Numbers o f taxa present in southern Africa No. families No. genera No. species Bryophyta Pteridophyta Gymnospermae Monocotyledonae Dicotyledonae Total 88 28 6 37 163 322 291 74 6 502 Í 794 2 667 821 251 43 4 491 15 881 21 487 Non-seed plants Vascular plants Seed plants Flowering plants 116 234 206 2<K) 365 2 376 2 302 2 2% 1 072 20 666 20 415 20 372 + No. infrasp, taxa _ Species & infrasp. taxa Unpubl. taxa _ Total species & infrasp. taxa 0 17 0 429 1 803 2 249 821 268 43 4 920 17 684 23 736 5 5 0 209 t21 340 826 273 43 5 129 17 805 24 076 17 2 249 2 232 2 232 1 089 22 915 22 647 22 604 10 335 330 330 1 099 23 2S0 22 977 22 934 TABLE 2. — Numbers of naturalized and indigenous families, genera and species in southern Africa No. No. No. families ■ - naturaliz. = indigcn. families families No. genera No. No. - naturaliz. = indigen. genera genera No. species No, Total No. - naturaliz. + unpubl. = indigenous species species species Bryophyta Pteridophyta Gymnospermae Monocotyledonae Dicotyledonac Total 88 28 6 37 163 322 0 1 1 1 8 11 88 27 5 36 155 311 291 74 6 502 1 794 2 667 0 4 1 36 189 230 291 70 5 466 1 605 2 437 821 25! 43 4 491 15 881 2! 487 0 9 6 119 514 648 5 5 0 183 21 214 826 247 37 4 555 15 388 21 053 Non-seedplants Vascular plants Seedplants Flowering plants 116 234 206 200 ! 11 10 9 115 223 196 191 365 2 376 2 302 2 296 4 230 226 225 361 2 146 2 076 2 071 1 072 20 666 20 415 20 372 9 648 639 633 10 209 204 204 I 073 20 227 19 980 19 943 B othalia 15, 3 & 4 (1985) 615 com pare th e cu rrent List o f Species to th e earlier es tim ates of Oyer (1975, 1976) and G oldblatt (1978), w ho published only species counts. Previous estim ates and counts for the num ber o f indigenous species in Africa have been considerably sm aller than the totals presented in the List o f Species. F o r seed plants, Killick (1971) estim ated 17 500 species, D yer (1975, 1976) estim ated about 18 400 species, and G oldblatt (1978) reported 18 532 species, w hereas the count obtained from the List o f Species (1984) is 19 980. A hand count o f the in digenous species o f vascular plants m ade by W ells et al. (1983) yielded 20 044 species, which is w ithin 100 species o f th e count o f 20 139 determ ined from the List o f Species. This is independent confirm ation o f th e accuracy o f the com puterization process, The count was o b tained by different individuals using a different m ethod for a different purpose, but w ork ing on th e sam e herbarium collection at the sam e tim e. flora a re shown in Table 3. All these families are flowering plants, and account for over 82% o f the entire flora, and 87% o f th e flowering plants. Tw elve families have around 500 o r m ore species and infraspecific taxa, with a break o f nearly 100 taxa betw een the rest, which have few er than 400 taxa. T he 12 largest families account fo r over 58% of the total flora and over 60% o f the flowering plants. A s currently treated, the largest family is M esembryanthem aceae, but it is believed th at critical re vision will bring the num ber o f accepted taxa down to ab o u t 1 200 (G ibbs Russell & G len , 1984). T h e family would then rank third, below A steraceae and Fabaceae. The largest families in the southern African flora T hese 38 families are only 12% o f th e to tal num b er o f fam ilies, and the 8 families that contain about half the total flora are less than 3% o f the total num b er o f families. W hen the families are arranged according to the num ber o f species they contain, it is found th at o ver half of all families have few er than 8 species and infraspecific taxa. T h e 38 families th at com prise m ore than 100 species a n d constitute 0,5% o r m ore o f the total G ood (1974) lists the 30 largest families in th e w orld. T en o f the 12 largest families in southern TA B LE 3, — The 38 families o f flowering plants with m ore than 100 species ranked by numbers o f species and subspecific taxa Rank 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 Family No. spp. & infrasp. taxa No. spp. No, genera Spp/, gen. % of total flora Running total of spp. & infrasp. taxa Running % of flora 2 684 2 417 1 802 1 142 1 024 984 955 892 568 526 487 480 392 362 346 306 301 279 273 257 243 236 233 22! 218 210 205 203 199 186 181 180 178 171 161 150 130 120 2 408 2 116 I 540 1 066 858 804 783 769 543 461 464 439 366 351 215 291 282 267 256 225 195 207 184 176 214 205 198 159 189 175 141 150 176 113 139 129 104 106 123 180+50* 122+13* 57+2* 46+1* 24 171 + 28* 58+1* 53+5* 43+5* 36 54 13+1* 41 5 20+1* 12 5 12 32+5* 2 59+1* 21 35+7* 10 4 17 8 9 7 6 13+4* 6 15 + 19* 15+6* 13+2* 14+1* 10+4* 19,5 11,7 12,6 18,7 18,6 33,5 4,6 13,3 10,2 10,7 12,9 8,2 28,2 8,6 43,0 14,6 23,5 53,4 21.3 7,0 97,5 3,5 8,8 5,0 21,4 51.3 11,6 19.9 21.1 25,0 23.5 11.5 29,3 7,5 9,3 9,9 7.4 10,6 11.2 10,1 7,5 4,8 4.3 4,1 4,0 3.7 2,3 2,2 2,0 2,0 1,6 1,5 1,0 1,3 1,3 1,2 1.1 1,1 1,0 1,0 1,0 0,9 0,9 0,9 0,9 0,9 0.8 0,7 0.7 0.7 0.7 0,7 0,7 0.6 0,5 0,5 2 684 5 101 6 903 8 045 9 069 10 053 11 008 11900 12 468 12 994 13 481 13 961 14 353 14 715 15 061 15 367 15 668 15 947 16 220 16 447 16 720 16 956 17 189 !7 410 17 628 17 838 18 043 18 246 18 445 18 631 18 812 18 992 19 170 19 341 19 502 19 652 19 782 19 902 11,2 21,3 28,8 33,5 37,8 41,9 45,9 49,6 52,6 54,1 56,2 58,2 59,8 61,3 62,8 64,0 65,3 66,5 67,7 68,6 69,6 70,6 71,6 72,5 73,4 74,3 75,2 76,1 76,9 77,6 78,4 79,1 79,9 80,6 81,3 81,9 82,4 82,9 Mcscmbryanthcmaceac Asteraceae Fabaceae Liliaceae Iridaceae Ericaceae Poaceae Asclepiadaceae <s.s.) Scrophulariaceae (s.s.) Euphorbiacea Cyperaceae Orchidaceae Proteaceae Acanthaceae Crassulaceae Rutaceae Restionaceae G eraniaceae Campanulaeeae (s.s.) Lamiaceae Oxalidaceae Rubiacea Aizoaceae Apiaceae Selaginaceae Poiygalaceae Amaryllidaceae Rhamnaceae Thymeiaeaceae Sterculiaceae Lobeliaccae Rosaceae Santalaceae Brassicaceac Malvaceae Chcnopodiaccae Convoivulaccae Anacardiaceae 2 Calculated from Goldblatt (1978) G ood (1974) 'N aturalized taxa % of Rank in endem, 30 largest spp.* world famsb 98,0 86,0 74,5 88,5 96,5 99,7 44,4 86,9 86,8 79,7 51,8 80,5 97,0 66,4 88,6 94,1 94,1 96,3 92,1 63,1 98,0 52,1 98,0 90,3 97,1 87,5 81,9 92,3 95,0 84,6 (92) 94,0 94,4 91,7 32,0 85,2 29,8 72,0 — 1 3 11 — 21 4 13 8 6 9 2 — 15 — — — — — 7 — 5 — 17 — — — — — — — 24 — — — — — — B othalia 15, 3 & 4 (1985) 616 Africa are included in G o o d ’s list, but M esem bryanthem aceae and Iridaceae a re not am ong the w orld’s 30 largest families. T he 38 most im portant southern A frican families include 18 o f the largest families listed by G ood. T h e species/genus ratios o f the families can reflect th eir phytogeographical affinities. T he overall species/genus ratio for southern A frican seed plants is ab o u t 9,6. T he families with a species/genus ratio m ore th an twice the overall ratio are those known to have diversified extensively within southern Africa especially in Capensis: M esem bryanthem aceae, Liliaceae, Iridaceae, E ricaceae, P roteaceae, Crassulaceae, R estionaceae, G eraniaceae, C om panulaceae, O xalidaceae, Selaginaceae, R ham naceae, Thym elaeaceae, Sterculiaceae, Lobeliaceae and Santalaceae. Fam ilies with a species/genus ratio about half th e overall ratio o f 9,6 a re all families o f worldwide distribution o r with centres o f diversity in the tropics o r n o rth tem p erate areas: Poaceae (w orldw ide), Rubiaceae (pantropical), and A piaceae (north tem p erate) (G o o d , 1974). All the families with a high species/genus ratio also have m ore than 88% o f th eir species endem ic to southern A frica. H ow ever, not all families with a high percentage o f endcm ic species have a high species/genus ratio. In these exceptions, eith er more than half the genera in the family are endem ic but each have a m o d erate num ber o f species (R utaceae, A piaceae), o r less than a fourth o f the genera in the family are endem ic but a few genera have a large num ber o f endem ic species (Brassicaceae — Heliophila, R osaceae — Cliffortia). In T able 3, the families a re ranked by the num ber o f species and subspecific taxa. If the num ber o f in digenous specics is used fo r ranking instead, there is little change in th e placing o f the 38 largest families. C yperaceae and Euphorbiaceae exchange places at ranks 10 and 11, but th ere is only a difference o f th ree species betw een them . C rassulaceae falls from 15th to 20th as a result o f the large num ber o f sub specific taxa now accepted, and A piaceae falls from 24th to 30th as a result o f the high num ber o f natura lized species. Polygalaceae and T hym elaeaceae each gain th ree places in com parative ranking, mainly b e cause they have few naturalized species o r infraspesific tax a, and therefore th ere is little difference be tw een the total num ber o f taxa and the total num ber o f indigenous species. The 21 families o f flowering plants with m ore than 20 genera are listed in T able 4. O nly o n e family, A m aranthaceae, does not also have m ore than 100 species. T he ranking o f the families by num ber of genera corresponds only roughly co the ranking by num ber o f species and infraspecific taxa, but th e 12 families with th e most genera include ten o f the larg est families ranked by species and infraspecific taxa. Ericaceae and C yperaceae have com paratively fewer genera, and Rubiaceae and A piaceae have com para tively m ore. T he families in T able 4 can be divided into three groups o n the basis o f the percentage o f their genera that occur in southern A frica related to their world wide distribution, (a) T h ree families have from nearly half to virtually all th eir genera represented in southern Africa: M esem bryanthem aceae, the most strongly ‘southern A frican' o f all o u r families; Irida ceae, concentrated in the southern hem isphere; and Ericaceae, with the subfamily E ricoideae present in E urope but concentrated in southern Africa (G ood, 1974). (b ) All the families w ith a worldwide distribu tion described by D yer (1975, 1976) as simply ‘cos m opolitan’ have betw een 20 and 40% o f their genera represented in southern A frica, (c) T he families with few er than 20% o f their genera occurring in south ern Africa are distributed predom inantly in climatic TA B LE 4. — The 21 families of flowering plants with more than 20 genera ranked by numbers o f genera Rank Family 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 Asteraceae Poaceae Fabaccac Mesembryanthemaceae Rubiaccac Liliaceae Asclepiadaceae Scrophulariaceae Orchidaccae Euphorbiaceae Iridaccac Apiaceae Acanthaceae Lamiaceae Cyperaceae Brassicaceac Ericaceae Amaranthaceae Rutaceae Aizoaceae Malvaceae * D yer (1975, 1976) ’ Naturalized genera No. genera in sthn Africa 180+50' 171+28* 122+13' 123 59+1* 57+2* 58+1* 53+5* 54 43+5* 46+1* 35+7* 41 32+5* 36 15+19* 24 22+2* 20+1* 21 15+6' No. genera % of genera worldwide' indigenous in sthn Afr, 900 660 600 124 500 230 200 170 725 300 75 260 250 170 90 370 50 65 150 •> 75 20,0 25,9 20,0 99,2 11.8 24,7 29,0 31.1 7.4 14.3 61.3 13.5 16,4 18,8 40.0 4,1 48,0 33,8 13,3 — 20.0 Family rank in sthn Afr, by spp. & infrasp. taxa Worldwide family distribution' 2 7 3 1 22 4 8 9 12 10 5 24 14 19 11 34 6 Cosmopolitan Cosmopolitan Cosmopolitan Southern Africa C os., tropical Cosmopolitan C os., subtropical Cosmopolitan C o s., absent dry area C os., tropical C os., sthn hemisphC os.. tem perate Trop, & subtrop. C os., warm temp. Cosmopolitan C o s., n o n h em temp. C o s.. Europe & sthn Afr. C os., trop. & warm C o s., warm & temp. Cosmopolitan C os., trop. & subtrop. — 15 23 35 617 B othalia 15, 3 & 4 (1985) TABLE 5 . — The 35 genera o f flowering plants with m ore than 100 species and infraspecific taxa Rank 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 Genus Erica Ruschia Conophytum Aspalathus Senecio Euphorbia Helichrysum Oxalis Lampranthus Indigofera Pelargonium Phylica Delosperma Aloe Thesium Haworthia Hermannia Gladiolus Agathosma Crassula Wahlenbergia Cliffortia Sutera Selago Restio Drosanthemum Muraltia Stapelia Schizoglossum Lotononis Cheiridopsis Sphalmanthus Lobelia Euryops Disa No. supp. & infrasp. taxa No. indigen. spp. Family 792 372 362 333 323 302 260 241 237 226 219 190 180 175 168 165 155 148 138 137 136 129 128 128 119 119 119 119 117 114 111 111 103 103 101 638 352 301 256 309 266 241 193 218 212 211 147 159 152 166 153 146 107 138 42 135 108 125 127 113 111 116 88 97 104 103 108 81 90 90 Ericaceae Mesembryanthemaceae Mesembryanthemaceae Fabaceae Asteraceae Euphorbiaceae Asteraceae Oxalidaceae Mesembryanthemaceae Fabaceae Gcraniaceac Rhamnaceae Mesembryanthemaceae Liltaceac Santalaceae Liliaceae Sterculiaceae Iridaceae Rutaceae Crassulaceae Campanulaceae Rosaceae Scrophulariaceae Sclaginaccac Restionaceae Mesembryanthemaceae Polygalaceae Asclepiadaceae Asclepiadaceae Fabaceae Mesembryanthemaceae Mesembryanthemaceae Lobeliaccae Asteraceae Orchidaceae areas not well represented in southern Africa: Orchidaceae (absent from dry areas), Brassicaceae (n o rth tem p erate), Euphorbiaccae (tropical), Apiaccae (tem p erate), A canthaceae (tropical and subtro pical). A m aranthaceae and R utaceae are excep tions. B rassicaceae is notew orthy because over half its genera in southern Africa are naturalized. Tw o bryophyte families also have m ore than 20 gen era, P ottiaceae (29) and L ejeuneaceae (28). The largest genera in the southern African flora Thirty-five genera in the southern A frican flora have m ore th an 100 species and infraspecific tax a, as shown in Table 5. All o f these genera occur in one o f the 38 largest families, and 22 occur in o ne o f th e 12 families with o ver 500 species and subspccific taxa. A m ong th e 12 largest families only tw o, Poaceae and C yperaceae, d o not have a genus with m ore th a n 100 species. A ccording to G oldblatt (1978), only five of th e large genera are endcm ic, three in M esem bryan them aceae (Ruschia, Conophytum and Lampranthus), one in F abaceae (Aspalathus) and one in R u taceae (Agathosma). N ot surprisingly, the 12 largest families have the most large genera. M esem bryanthem aceae has 7 genera with o v er 100 taxa, A steraceae and Fabaceae have 3 each , and Liliaceae and A sclepiadaceae 2 Family rank in sthn Afr. by spp. & infrasp. taxa 6 1 1 3 2 10 2 21 I 3 17 28 1 4 33 4 30 5 15 20 18 32 9 25 16 I 26 8 8 3 1 I 31 1 12 each. T he o th er families each have only a single genus with m ore than 100 taxa. Size and composition o f the southern African flora compared to the floras o f other parts o f Africa Since 1950, several com plete accounts o f th e flora o f various parts o f Africa have b een published, cov ering the areas shown in Fig. 1. T h e num bers of taxa, num ber o f vegetation types, cen tres o f en d e m ism , and sizes o f each o f these areas can be co m pared to southern A frica, in o rd e r to place o u r flora into perspective with that o f o th e r parts o f the conti nent, and with sm aller areas inside southern A frica. T he treatm ents considered fo r tropical A frica w ere: Flora o f West Tropical Africa, 2nd edn (K eay, 1954, 1958; H epper, 1963, 1968, 1972); The flowering plants o f the Anglo-Egyptian Sudan (A ndrew s 1950, 1952, 1956) and Enumeratio Plantarum Aethiopicae Spermatophyta (C ufodontis, 1953-1970, as listed by M eyer, 1973). F or areas w ithin southern Africa, the following treatm ents were considered: Prodromus einer Flora von Sudwestafrika (M e rx m iiller, 1966-1970); Flora o f Natal (R oss, 1972); Flora o f Lesotho (Jacot G uillarm od, 1971); Flora o f Swazi land (C om pton, 1966, 1976); Plants o f the Cape Flora (G oldblatt & B ond, 1984) and Flora o f the Cape Peninsula (A dam son & S alter, 1950). 618 B othalia 15, 3 & 4 (1985) w « kt *r vegetation types and num ber of centres of ende mism. Southern Africa has the largest num ber o f fa milies, genera, and species plus infraspecific taxa. A lthough it does not cover the largest a re a , it has the largest num ber o f vegetation types and centres of endem ism as m apped by W hite (1983). A lthough the west tropical A frican flora covers nearly twice the area o f southern A frica, it has less than a third o f the num ber o f species and infraspecific taxa as southern Africa. T he richness o f the southern A frican flora is further em phasized w hen areas within it are com p ared to areas outside. T he included C ape flora ranks second in num ber o f species, with at least 1 500 taxa m ore than the third-ranking west tropical African flora, even though the area covered by the west tropical A frican flora is m ore than 53 tim es the area o f the C ape flora. T he included South W est A frican/N am ibian flora is sim ilar to the Sudan flora in num ber o f species and infraspecific tax a, even though S udan has m ore than twice the num ber of vegetation types and nearly twice the area o f South W est Africa/Namibia. T he far larger num ber o f taxa recorded for south ern A frica may be ascribed to four factors; FIG- 1.—Parts o f Africa covered by floras considered in this study. 1. Flora of West Tropical Africa; 2, the Flowering Plants o f the Sudan; 3, Enumeratio Plantarum Aethiopicae Sperm atophyta; 4, List o f Species o f Southern African Plants; 5. the Flora o f Swaziland; 6 , Flora o f N atal; 7, Flora o f Lesotho; 8. Plants o f the Cape Flora; 9 , Flora o f the Cape Peninsula; 10. Prodromus einer Flora von Siidwestafrika. T ables 6 ,7 & 8 com pare num bers o f fam ilies, g en e ra and species plus infraspecific taxa for each o f the te n floras, and Table 9 shows each o f the floras ranked by num bers o f taxa, area covered, num ber of 1. southern A frica has the largest num ber o f vege tation types o f all the floras considered. N one o f the vegetation types are shared betw een southern Africa and the tropical floras north o f the eq u ato r except the habitat-lim ited and com paratively small aquatic, halophytic, m angrove and afrom ontane vegetation types. T he southern A frican vegetation includes the C ape flora, with over 8 500 species and the karroid vegetation types, also rich in species, 2. southern Africa has four centres of endemism (W hite, 1983) and a very high percentage o f endem ic taxa (G oldblatt, 1978). Tw o o f the centres o f en d e T A B LE 6.— Comparison o f numbers o f families in African floras Tropical floras N. of equator W trop Afr. Sudan Gymnosperms Monocotyledons Dicotyledons Total seed plants 3 40 159 202 4 32 135 171 Pteridophytes Total vascular plants 27 229 Included sthn African floras Entire sthn Ethiopia Afr. flora Swaziland Natal Lesotho Cape flora Cape Penins. SWA/ Namibia 3 32 143 178 6 37 163 20« 2 2! Ill 134 4 34 141 179 0 17 78 95 4 32 116 152 3 19 96 118 1 26 127 154 — .— 13 147 18 113 — — 28 234 — — 11 129 12 166 — — TA B LE 7,— Comparison o f numbers of genera in African floras Tropical floras N, o f equator Included sthn African floras en tire sthn Ethiopia Afr. flora Swaziland W trop Afr. Sudan Gymnosperms Monocotyledons Dicotyledons T otal seed plants 3 c.368 1 298 I 669 4 255 878 1 137 3 296 1 090 1 389 6 502 1 794 2 302 Pteridophytes T otal vascular plants 72 1 741 — — — — 74 2 376 Natal Lesotho Cape flora Cape Penins. SWA/ Namibia 2 200 558 760 4 311 918 1 233 0 159 367 526 4 282 700 986 3 209 459 671 1 205 481 687 36 796 — — 28 554 28 699 16 703 — — B othalia 15, 3 & 4 (1985) 619 TA BLE 8.— C om parison o f n u m b ers o f species and infra specific taxa in A frican floras T ro p ical floras N. o f e q u a to r W tro p Afr. Sudan G ym nosperm s M onocotyledons D icotyledons T o ta l se e d plants 4 1 810 5 200 7 014 4 817 2 393 3 215 Pteridophytes T o tal vascular plants 312 7 326 Included sthn A frican floras E ntire sthn E thiopia Afr. flora Sw aziland N atal L esotho C ap e flora* Cape Penins. SWA.' Nam ibia 4 1 527 4 792 6 323 43 S 129 17 80S 22 977 8 566 1 515 2 089 14 1 304 3 500 4 818 0 485 1 052 1 537 11 2 075 6 418 8 504 6 846 1 713 2 565 1 740 2 384 3 125 __ _ — 273 23 250 73 2 162 _ — 54 1 591 c.75 8 579 57 2 622 43 3 168 — 'S p e c ie s only. B ond & G o ldblatt (1984) did not include infraspecific taxa. T A B L E 9. — R ankingof floras by num bers o f taxa o f se ed plan ts .a r e a , num ber o f vegetation types and n u m b er o f c e n tre s o f endem ism R an k Fam ilies G enera Spp. & infrasp. taxa A pprox. area (ktn: ) N o. vegetation types-' N o . centres o f endemism*1 S thn Afr. 206 Sthn Afr. 2 302 Sthn A fr, 22 977 W . tro p , Afr, 4 836 700 Sthn A fr. 28 S th n Afr, 4 2 W trop. Afr. 202 W trop. Afr. 1 669 C ape flora 8 504" S thn Afr. 2 573 000 W tro p . A fr. 24 W tro p . Afr. 3'. 3 Natal E thiopia I 389 W tro p . Afr. 7 014 Sudan 2 505 800 S udan 18 E thiopia 3 178 N atal I 233 E thiopia 6 323 E thiopia 1 696 450 E thiopia 17 SW A 2 5 Sudan 171 Sudan 1 137 N atal 4 818 SW A 825 200 SW A 10 C ap e flora 2 6 SW A 154 C ape flora 986 S udan 3 215 C ape flora 90 000 N atal 7 Sudan 2 7 C ap e flora 152 Swaziland 760 SW A 3 125 N atal 86 000 C ape flora 4 Natal 1 8 Swaziland 134 SW A 6R7 C ap e Penins. 2 565 L esotho 30 276 L esotho 4 L esotho 1 9 C ap e Penins. 118 C ape Penins. 671 Swaziland 2 089 Swaziland 17 363 Swaziland 3 C ap e Penins. 1 10 L esotho 95 L esotho 526 L esotho 1 537 C ape Penins. 470 C ape Penins. 1 Sw aziland 0 1 179 4 E thiopia * W hite (1983). ‘ Species o nly. B ond & G oldblatt (1984) did not include infraspecific taxa. m ism , th e C ape and the K aroo-Nam ib, occur only within southern A frica (except for a small extension o f th e K aroo-N am ib into southern A ngola), and al though the Z am bezian centre lies mostly outside southern A frica, it is not covered by any o f the Flo ras considered except the List o f Species o f Southern African Plants. Only the afrom ontane centre of endemism is shared with other Floras considered. G oldblatt (1978) estim ates th at 80% o f the southern A frican species are endem ic, and the levels o f e n d e mism in m any o f the largest families are considerably higher than 80% , as shown in Table 2. B rcnan (1978) calculated an area/endem ic index fo r various parts o f tropical Africa. T his index estim ated for the w hole o f southern Africa is 161 (i.e, 20 000 x 0,8 = 16 000 estim ated endem ic seed plant species; 2 573 000 km ; -s- 16 000 endem ic species = 161 area/endem ic). This indicates higher levels o f e n d e mism fo r southern A frica than fo r anywhere in tropi cal A frica, w here Brenan reports strongest e n d e mism in G abon (239) and C abinda (251). In com parison, he reports that Sudan, w ith a sim ilar area to southern A frica, has an area/endem ic index o f 50 000. A t the o th er extrem e, the area/endem ic in dex calculated for the C ape flora using the figures o f B ond & G oldblatt (1984) is 15, 3. the counts fo r the southern A frican flora as a whole and fo r the C ape flora are o f m ore recent date than any o f th e others. M any m ore taxa are probably known fo r each o f the areas now than ap p ear in the Floras considered, m ost o f which a re o v er ten years old. F or exam ple, the num bers o f seed plant taxa reported by Jacot G uillarm od (1971) fo r L esotho w ere 526 genera and 1 591 species and infraspecific taxa. A listing from PREC IS o f all L esotho speci m ens m ade in 1984 show ed 702 genera and 2 726 species and infraspecific tax a, an increase o f 25% and 42% respectively in 13 years. Sim ilarly, W hite (1983) estim ates 8 000 species for the G uineo-Con- 620 B othalia 15, 3 & 4 (1985) golan cen tre o f endem ism , which form s only a part o f th e area o f the west tropical A frican flora, result o f the sam e factors th at give th e Poaceae a com paratively low ranking in the sam e floras. 4. the southern A frican flora is being studied pri marily by botanists w orking in the region, w hereas th e floras o f west tropical A frica, Sudan and E thio pia have been studied prim arily by botanists working in E urope. This undoubtedly increases the p ro p o r tion o f the existing flora th a t has been collected. Even within southern A frica, m apping the num bers o f specim ens per q u arte r degree square has shown that collecting intensity is greatest in easily acces sible locations (G ibbs R ussell, R etief & Sm ook, 1984), T hree families show differences betw een the m a jor com ponents o f the tropical and southern African floras, A canthaceae and Rubiaceae are am ong the ten largest families both in the tropical floras north o f the eq u ato r and in the included floras o f tropical affinity within southern A frica, nam ely South West A frica/N am ibia, N alal and Swaziland, Asclepiada ceae is the only family th at is a m ajor com ponent o f all (extra-C apensis) southern A frican floras but not o f the tropical floras north o f the equator. T h e individual floras included in the entire south ern African area each has only a fraction o f the total num ber o f taxa in the region, and only a small pro portion o f th e 28 vegetation types present in the whole area. N evertheless, only the th ree karroid ve getation types are not covered by at least one o f the included Floras considered. T h e ten largest families in each o f the Floras are com pared in T able 10. All the ten largest families in the en tire southern A frican flora are also am ong the largest families in at least one o th er F lora consid ered. Each o f th e 24 families concerned is represen ted in southern A frica, and only A pocynaceae and A nnonaceae d o not have o v er 100 species and infraspecific taxa in southern Africa, A m ong the floras considered, these tw o families are a m ajor com po nent only o f th e west tropical A frican flora. Only tw o families, A steraceae and Fabaceae are am ong the ten largest in all the floras. A steraceae ranks first o r second in all the included southern A frican floras, and if M esem bryanthem aceae were critically revised (G ibbs Russell & G len , 1984) A s teraceae would be the largest family in the entire southern A frican flora as well. In the tropical floras A steraceae is not as outstanding, ranking below Fa baceae and Poaceae. In contrast, F abaceae is o f first o r second rank in the tropical floras, but ranks only from second to fourth in the southern A frican floras. Poaceae is am ong the ten largest families in all the floras considered except the C ape flora, and it has a much lower ranking in southern A frica as a whole th an in th e tropical floras o r in the included floras o f southern Africa, N evertheless, in absolute num bers there are m ore taxa recorded fo r Poaceae in south ern A frica than fo r the tropical floras even though the ranking in southern Africa is lower. This com paratively low er rank for southern A frica as a whole may result from two factors. Firstly, the many w ide spread grass species are counted separately for the sm aller included floras, but only once for the entire southern A frican flora. Secondly, th e C ape flora has a very low num ber o f grasses in com parison to Aizoaceae (s.I.), Liliaceae (s.l.), Iridaceae and Erica ceae. T h e large num bers o f taxa in these families in the C ape flora give the Poaceae a lower ranking in th e southern A frican flora as a whole, C yperaceae is th e only family that is a m ajor com ponent o f all flo ras considered except the entire southern African flora and the C ape flora. Its absence from the ten largest families in these tw o floras is probably the Lam iaceae shows a difference betw een eastern Africa and the rest o f the continent. In A frica, this family has its greatest im portance in eastern Africa, from Sudan and Ethiopia south to Swaziland and N atal (w here it ranks 11th). The families th at are peculiarly southern A frican, recognized in T able 3 by th eir high species/genus ra tios and percentage o f endem ism , are also clearly shown in T able 10. A izoaceae, E ricaceae, Iridaceae and R estionaceae are am ong the ten largest families in the C ape flora, and through it, o f southern Africa as a whole. Proteaceae and R utaceae are am ong the ten largest families only in the C ape flora. Am arvllidaceae and C am panulaceae a re m ajor com ponents only o f the high-altitude L esotho flora, although C am panulaceae ranks 11th in the C ape flora. Again, one family exhibits an opposite p attern: E upho rb ia ceae is one o f the ten largest families in all floras ex cept the C ape and Lesotho. Convolvulaceae and M alvaceac arc m ajor com po nents of the flora only in Sudan, although they have m ore than 100 species in southern Africa. T hree families show patterns o f distribution that do not coincide with any others. Liliaceae (s.l.) is one o f the ten largest families in southern and east ern A frica, but not o f west tropical Africa and Su dan. O rchidaceae, although the sccond largest fam ily in the w orld, is not am ong the m ajor com ponents of the floras o f South W est A frica/N am ibia, Sudan o r E thiopia, perhaps because o f the considerable arid areas in these countries. In C apensis, O rchida ceae ranks 12th in the C ape flora as a w hole and 7th in the C ape Peninsula. Scrophulariaceae is the most unusual in its areas o f im portance. It is am ong the ten largest families in the C ape flora (tem perate and w inter rainfall), South W est Africa/Nam ibia and Su dan (both tropical and a rid ). N atal (o f tropical affin ity and mesic to arid), and L esotho (high altitude). Species richness o f the souihern African flora com pared to floras o f other parts o f the world The richness o f the southern A frican flora com pared to floras o f o th er large regions o f the world, both tropical and tem perate, is shown in T able 11. T he species/area ratio fo r the whole o f tropical Africa is sim ilar to that o f its included parts, Sudan and west tropical Africa. T he ratio for southern Africa is ab o u t five tim es as great, illustrating the com parative poverty o f the tropical African flora discussed in detail by B renan (1978). Tw o o th er tropical areas, Brazil in tropical S outh A m erica and tropical A sia, are widely separated geographically B othalia 15, 3 & 4 (1985) 621 T A B L E 10. — C om parison o f n um bers o f species and infraspecific taxa in th e 10 largest families in all floras considered. The top num ber is the rank o f the family in the flora and the b o tto m n u m b er is the n u m b er o f species and infraspeciftc taxa. A d a sh in the top position shows that th e family is n o t am ong th e ten largest in that flora Included sthn A frican floras T ropical floras N o f e q u a to r W trop. A fr. Sudan E thiopia slh n Afr. flora 8 178 7 98 4 273 — 0 — 0 — (25) 1 2 917 --(17) — (19) __ (205) 10 112 — (15) __ (13) A pocvnaceae 9 132 A sclcpiadaccac (s./.) A canthaccac A izoaceae (s./.) A m aryllidaeeae A nnonaceae —(14) (362) — (27) __ 03) Swazi land N atal L esotho C ap e flora3 C ape Penins. SW A ' N am ibia 10 64 _ __ (108) __ (27) __ 0 7 118 — <9) — (39) — (6) --(54) (6) --(18) 2 769 9 94 4 191 10 34 --(83) — (27) — (2) __ (6) --0 --(0) — 0 — (25) — 0 — (6) — (!7 ) — — (23) (8) — (33) — (41) — (12) — (94) — (59) — (150) 8 915 5 96 5 207 7 60 — <135) — (I) — (21) A steraccae 6 291 3 208 3 565 2 2 417 1 223 1 546 I 302 1 986 I 292 2 34? C am panulaceae (s.l.) — (27) — (16) — (36) —(13) (454) — (21) — (75) 9 35 — (222) — (68) — (26) C onvolvulaceae --(76) 8 77 __ (85) —(37) (130) — (22) — (52) (13) 5 312 4 152 7 232 —(11) (487) 7 71 7 168 — (3) — — (3) (7) — (9) — (32) 7 272 6 131 6 235 10 506 1 725 I 390 2 659 3 1 802 — (9) —. (27) 1 024 9 74 5 242 — (57) 8 220 — (78) — (71) -> 10 152 612 387 C yperaceae E ricaccac E uphorbiaceae F abaceae (s.l.) Iridaceae L am iaceae L iliaceae [s.l.) M alvaceae __ (15) —, (% ) — (65) Poaceae 1 704 6 122 — (15) 3 672 8 119 — 0 — 420 4 644 2 191 1 264 __ (36) — (92) 10 34 5 612 4 154 — (23) 9 68 __ (U 7 ) __ (26) .— (44) — (15) — (47) 4 122 4 226 5 83 6 419 6 123 5 139 — (35) (161) — (28) — (60) — (13) — (27) — (13) — (56) 7 3 173 3 410 2 170 — (181) 3 181 I 344 -(1 3 ) 392 — m — (16) — (4) 7 320 — (47) -—(12) (480) 6 77 6 206 8 54 — (206) 7 121 (0 — (9 ) — — 8 310 10 86 — 0 — (54) — (16) — (41) 10 259 — (23) — (3) — (60) (100) 5 — 119> (257) 4 1 142 955 — (49) R estionaceae — 0 — 0 — 0 —<16! (301) R ubiaceae 3 557 5 152 9 204 R utaccae — (28) — (9) — (23) Scrophulariaceac — (93) 1(1 64 — (133) 51 5 133 10 89 4 403 • c/c o f to ta l flora (seed p lan ts) in 10 largest families — (44) — (?35) O rchidaceae T o tal no. sp p . & infrasp. taxa 3 594 in 10 largest families — (9) 6 83 (IS ) — (203) — (98) — (3) (!) — (123) — 8 114 (I? ) 4 94 — (6) Protcaccae — 6 984 — (->> — (10) 8 70 2 179 8 146 2 — (6) —(22) (236) (!) — (62) 9 130 (1) — (18) —(16) 306 — (8) — (13) — (2) — (53) 10 127 3 121 9 310 9 786 — 9 1 733 3 486 13 468 1 143 2 586 1 070 5 301 I 494 I 822 54 55 59 55 54 69 62 58 58 “Spccies only. B ond & G oldblatc (1984) did not include subspecific taxa. B othalia 15, 3 & 4 (1985) 622 T A B L E 11. — Species/area ratios for several rceions o f the w orld w ith area s o f over ± 2 500 « 0« km ; R egion T ropical A frica W est tropical Africa Sudan Brazil India, P akistan, B angaladesh. Burm a A ustralia E astern N orth A m erica S outhern Africa S outhern Africa (excluding C apensis) N o. species A rea (km -) Species/A rea 30 000» 7 300 3 200 40 000“ 20 (X10 000 4 500 000 2 505 800 8 456 500 0.0015 0,0016 0.0013 0,0047 20 000" 25 OOO4 42Sd 20 000 4 7 3 2 885 280 716 000 238 000 573 (XK) 0,0041 0.0032 0.0014 0.0078 14 200' 2 483 000 0.0057 aB renan (1978). bG o o d (1974). ‘Ride (1978) q u o ted in G oldblatt (1978). dG o ldblatt (1978). ‘C alculated from G ibbs Russell ei at. (1984) a n d B ond & G o ldblatt (1984), but have sim ilar species/area ratios. A lthough tropi cal areas are known to have high concentrations o f species, the species/area ratio fo r southern Africa is about 1,7 tim es greater than either. T he A ustralian flora is som etim es com pared to that o f southern Africa because both are southern hem isphere areas with tropical and tem perate vegetation elem ents and both have high levels o f endem ism . H ow ever, the species/area ratio for southern A frica is nearly 2,5 tim es that o f A ustralia. T he ratio for eastern N orth A m erica, in the north-tem perate zone, is also far b e low that o f southern Africa. Even if the extrem ely diverse C ape flora is elim inated from the determ ina tion o f th e species/area ratio for southern A frica, the ratio is nevertheless higher than that fo r tropical South A m erica o r Asia. Size and composition o f the southern African flora reported in the List o f Species compared to previous recent treatments T able 12 shows the num bers o f genera and species recorded for each family in the List o f Species (1984), D yer (1975, 1976) and G oldblatt (1978). Fa milies th at are trea ted differently in the three sources are shown both sensu stricto and sensu lato so that com parisons can be m ade. For seed plants, the count o f G oldblatt and the estim ate based on D yer agree within 50 species. T hey d a te from the sam e p erio d , but D yer’s w ork was carried out pri marily at P R E . w hereas G oldblatt’s count was d e te r mined from several herbaria, literature, and consul tation with experts in various groups. T h e closeness o f the final count confirm s b oth as being reasonable determ inations for th at time. T he List o f Species count is about 215 indigenous genera and 1 450 species higher than G o ld b latt’s count. T his discrepancy is the result o f different num bers o f species recorded in a num ber o f families, For 63 families (30% o f the num ber o f families of seed plants) th e List o f Species has a higher num ber o f species th an eith er o f the o th er counts, and 27 fa milies exceed D yer’s and G oldblatt’s counts by m ore th an 5 species. O ver a third o f the species are in Mesem bryanthem aceae, which has been previously m entioned as having in reality far fewer species than are presently recognized. T h ree o th e r families differ by m ore than 100 species, A sclepiadaceae, F abaceae and Liliaceae. In each o f these fam ilies, certain gen era are under revision but others are still in need o f treatm ent. G oldblatt’s count has only th ree families for which the highest num ber o f species is recorded, O rchidaceae, R estionaceae and R ubiaceae. All three o f these families have had recent changes in species num bers as a result o f taxonom ic revision. A lthough D yer’s estim ate has the lowest total num ber o f species, curiously there are 15 families for which his species counts are the highest. Tw o o f these fam ilies, A steraceae and C rassulaceae, differ from the List o f Species by m ore than 100 species. A num ber o f genera in A steraceae and the whole of Crassulaceae have been revised since 1975. Simi larly, Poaceae and Sterculiaceae are recorded with over 50 species m ore by D yer, but w ork done for the F T E A (C layton & R envoize, 1982) and a m ore accu rate determ ination o f naturalized species has re duced species num ber in Poaceae and revisionary work in Hermannia has reduced species num bers for Sterculiaceae, It appears therefore, th at differences in counts of species in a num ber o f families are due to revisions com pleted o r in progress betw een the tim e o f com pletion o f D yer, G oldblatt and the List o f Species. H ow ever, the substantially higher counts in the List o f Species for A piaceae, C henopodiaceae, C yperaceae, E uphorbiaceae, F abaceae, Liliaceae and Scrophulariaceae cannot be solely a ttrib u ted to th e result o f further study in these families. F urtherm ore, even though revisionary w ork has resulted in a lower count o f species in families such as Poaceac and C rassulaceae, and in such genera as Helichrysum, w here 283 taxa (260 species) w ere recognized at P R E in 1981, but only 260 taxa (241 species) w ere recognized after H illiard’s treatm ent o f the genus for the Flora o f Southern Africa, revision does not necessarily result in the recognition o f a sm aller num ber o f species. T he revision o f Asparagus cu r rently being com pleted for the Flora now recognizes tw o genera and 77 species w here previously one genus with 44 species was recognized (A . A. O berm eyer, pers. com m ,). C . H . Stirton (pers. com m .) predicts th at there may be ultim ately a total o f 2 000 B othalia 15, 3 & 4 (1985) 623 TABLE 12.— Comparison o f numbers o f taxa per fam ily. Families are listed alphabetically w ithin each g ro u p (B ry o p h y ta. Pteridophyta, Gymnospermae, Angiospcrmae). The difference between counts in the List o f S pecics (1984), D yer (1975, 1976) and G oldblatt (1978) is indicated when there is a difference o f m ore than 5 species. N aturalized fam ilies and genera are indicated by an asterisk. This table runs from p. to p. List o f Species (1984) Fam ily B ry o p h y ta A crobolbaceae A delanthaceae A m blystegiaceae A ndreaeaceae A neuraceae A ntheliaceae A nthocerotaceae A rchidiaceae A rnelliaceae A ulacom niaceae A ytoniaceae Bar tram iaceae Brachytheciaceae B ryaceae B ryobartram iaceae C alym peiaceae C alypogejaceae C ephaloziaceae CephalozieUaceae Cleveaceae C odoniaccae C ryphaeaceae Dicranaceae D itrichaceae Encalyptaceae E nto d o n taceae Ephem eraceae E rpodiaccae Eustichiaceae E xorm othecaceae Fabroniaceae F issidentaceae Fontinalaceae Funariaceae G igasperm aceae G rim m iaceae G ym nom itriaceae H ed wig iaceae Herbertaiceae H ookcriaceae H ylocom iaceae H ypnaceae Jubulaceae J ungerm an niacea e Lejcuneaceae L em bophyllaceac Lepicoleaceae L epidoziaceae Leskcaceae L eucodontaceae L ophocoleaceae Lunular iaceae M aichantiaceae M eteoriaceae Metzgeriaceae Mn iaceae M onocarpaceae N anobryaceae Neckeraceae O rthotrichaceac O xym itraceae Pallaviciniaceae Phyllogoniaceae Plagiochilaceae Plagiotheeiaceae G enera 3 2 6 1 1 1 2 1 1 1 4 6 5 10 1 4 1 2 1 I 1 2 14 7 1 4 1 2 1 1 6 1 1 6 3 3 3 4 1 9 1 5 1 7 28 1 1 9 4 3 3 1 2 S 1 1 1 1 Species & infrasp. taxa 4 3 8 4 9 1 7 11 3 I 12 17 13 43 1 10 5 5 13 1 8 2 42 14 2 12 3 4 1 3 12 30 1 13 3 8 4 4 1 12 2 20 15 21 64 I 3 20 9 Dyer (1975,76)» Indigen. species 4 3 8 4 9 1 7 11 3 1 12 17 13 43 1 10 5 5 13 1 8 2 42 14 2 12 3 4 1 3 12 30 1 13 3 8 4 4 1 12 2 20 15 21 64 1 1 20 9 4 4 12 1 6 12 1 6 5 5 13 1 1 13 1 1 1 ! 7 27 7 27 1 3 1 6 1 17 1 6 1 17 2 8 8 4 6 1 2 G enera Species G oldblatt (1978) G enera - - - - - - - - ■ Species • • - - - - - - - • ■ - * - • * D ifference 624 B othalia 15, 3 & 4 (1985) List o f Species (1 9 8 4 ) Fam ily Polytrichaceae Porellaceae Pottiaceae Prionodontaceae Pterobiyaeeae Ptychom itriaceae Racopilaceae Radulaceae R habdoweisiaceae R hachitheciaceae Rhizogoniaceae Ricciaceae Riellaceae Schist ochilaceae Seligeriaceae Sem atophyllaceae Sphaerocarpaceae Sphagnaceae Splachnaceae Taïgioniaceae Thuidiaceae T rachypodaceae W aidiaceae Pteridophyta A diantaceae Aspidiaceae Aspleniaceae A thyriaceae Azollaceae Blechnaceae C yatheaceae Davalliaceae D ennstaedtiaceae Equisetaceae Gleicheniaceae Gram m itidaceae Hym enophyllaceae Isoetaceae Lindsaeaceae Lom ariopsidaceae Lycopodiaceae M aiattiaceae Marsiliaceae Ophioglossaceae Osm undaceae Polypodtaceae Psilotaceae Salviniaceae* Schizaeaceae Selaginellaceae T helypteridaceae V ittariaceae G ym nosperm ae Cupressaceae Pinaceae* Podocarpaceae Stangeriaceae W elwitschiaceae Zamiaceae Angiospermae A eanthaceae Achariaceae Aizoaceae (S.i.) Aizoaceae (s.1.) (see M esem bryanthem aceae) A lism ataceae A m aranthaceae A m aryllidaceae Anacardiaceae G enera 4 1 29 1 4 2 1 1 1 1 2 1 1 1 1 1 1 6 2 1 9 10 2 5+1* 1 2 1 4 1 2 2 1 1 1 1 1 1 2 6+1* 1 1* 4 1 2+1* 1 1 1• 1 1 1 1 41 3 21 (1 4 4 ) 4+1» 22+2* 17 10+4* Species &. infrasp. taxa Indigen, species G enera Species - . - - . _ _ _ . . . . . . . _ _ . . . - G enera Species - 12 5 73 1 6 9 2 9 2 1 2 37 S 2 1 11 1 9 1 5 9 2 1 12 5 73 1 6 9 2 9 2 1 2 37 5 2 1 11 1 9 1 5 9 2 1 63 22 32 6 3 10 2 6 6 1 3 2 11 11 1 9 10 1 16 8 2 15 1 1 7 6 12 1 52 22 29 5 2 7 2 4 6 1 3 2 11 11 1 9 9 1 16 8 2 13 1 0 7 6 11 1 3 2 4 1 I 28 3 0 4 1 1 28 1 1 1 1 1 1 3 0 4 1 1 30 351 3 184 2592) 43 3 22 (144) 353 4 172 (2 4 7 6 ) 6 55 198 105 4 23 16 13 4 52 221 93 362 3 233 (2 917) 6 81 205 120 G oldblatt (1978) Dyer (1 9 7 5 ,7 6 )a _ . . - _ - . . . . . . , - * . . . - . . . . . . - . . . « - - - . D ifference . _ . - I . 3 1 1 1 1 4 1 1 25 41 3 355 3 135 2021 3 20 17 10 3 53 199 93 . SL (+571) D (+23) SL (+12) B oth alia 15, 3 & 4 (1985) 625 List o f Species (1984) Fam ily A nnonaceae A piaceae A pocynaceae A ponogetonaccae A quifoliaccae A raceae Araliaceae A iecaceae A ristolochiaceae A sclepiadaceae (s.s) Asclepiadaceae (s.1. ) (see Pcriplocaceae) A stcraceac B alanitaceae B alanophoraceae Balsam inaceae Basellaceae B egonia ceae Bignoniaceae B om ba caceac B oraginaceae Brassicaceac Brunlaceae B urm anniaceae Burscraceae B utom aceae Buxaceae C abom baceae C actaceae C allitrichaceae C am panulaceae (s.s.) C am panulaceae ( i f .) (see L obeliaceae) C aneliaceae C annabaceae* C annaceae* C appaiaceae C aryophyllaceae C asuartnaceae* C elastraccac C eratophyllaceae* C henopodiaceae C lusiaceae (s.s.) C lusiaceae (s.l.) (see Hypericaceae) C om bretaceae C om m elinaceae C onnaraccae Convolvulaccae C ornaceae C rassulaccae C urcurbitaceae C unoniaceac C yperaceae Dicbapetalaceae Dioscoreaceae Dipsacaceae Droseraceae (s.s.) Droseraceae (s./.) (sec R oridulaccae) E benaceae E latinaceae Ericaceae E riocaulaceae E rythroxylaceae Escalloniaceae E uphorbiaceae F abaceae F lacourtiaceae Flagellariaceae Frankeniaceae Fum aria ceae D yer (1 9 7 5 .7 6 )a Species & infrasp. laxa Indigen, species G enera Species 13 221 41 9 1 18 11 6 2 892 (9 1 5 ) 13 176 38 7 1 16 11 6 1 769 (7 9 0 ) 7 37 17 1 1 5 3 5 1 54 (6 1 ) 14 142 39 5 1 12 10 6 2 700 (725) 180+50* 1 2 1 1+1* 1 6 1 13+3* 15+19* 12 1 1 1 1 1 1+3* 1 12 18 2417 4 4 S 2 6 11 1 105 171 78 1 33 1 2 1 13 3 273 454 2116 4 4 S 1 6 11 1 92 113 77 1 33 1 2 1 1 3 256 397 206 1 2 1 2 1 6 1 15 33 12 1 1 1 1 3 1 13 18 2 335 3 4 4 1 8 11 1 85 117 76 1 20 1 1* 1* 8 8+ f* 1* 12 1* 13+2* 2 1 1 2 63 66 1 59 1 150 11 1 0 0 50 39 0 56 0 129 9 1 1 1 0 5 7+1* 2 14+1« I 5 18 2 36 2 1 2 3 49 41 2 130 1 246 77 2 487 3 22 23 G enera 8 35+7* 16+2« 1 1 6 3 5 1 58+1* (66+1*) 2 2 24 2 2 1 43+5* 122+13* 13 1 1 4*1. G enera 8 32 16 1 1 5 3 5 1 60 174 1 2 1 1 1 1 13 15 12 1 1 - 2 1 1 3 257 397 1 1 1 1 18 1 Species 13 155 39 7 1 13 11 6 1 605 2072 3 3 4 1 5 11 1 81 109 75 1 31 2 1 1 2 380 12 12 1 2 1 49 40 58 108 2 8 - 2 47 47 0 64 1 80 10 21 41 36 2 107 1 115 72 2 464 3 20 23 1 21 5 8 2 15 ! 9 17 2 33 2 1 2 2 3 35 2S 2 107 1 261 69 2 385 4 18 18 19 21 5 6 2 15 1 5 17 2 30 2 1 2 2 3 35 28 2 94 1 219 67 2 421 3 17 24 19 21 51 12 984 14 6 1 526 1 802 26 1 3 8 35 11 804 13 6 1 461 1540 26 1 3 7 2 2 24 2 2 1 48 133 13 1 1 S 40 12 792 11 5 1 384 1 378 28 1 3 6 2 2 24 2 2 1 41 115 13 1 1 - 34 11 799 11 5 1 389 1 495 25 1 3 - - - - G o ld b latt (1978) 8 17 1 12 1 14 - 8 8 . Diffcrcnce SL (+34) SL (+185) D (+219) SL (+11) S L (+ 1 7 ) D (+ S ) D (+8) SL (+49) S L (+ « ) SL (+11) SL (+13) D (+146) SL (+79) SL (+77) SL (+162) 626 Bothalia 15, 3 & 4 (1985) List o f S pecies (1984) Fam ily Geissolom aceae G entianaceae G eraniaceae G esneriaceae G oodeniaceae Greyiaceae G rubbiaceae G unneraceae H aem odoraceae Haloragaceae (s.S.) Haloragaceae (s. 1.) (see G unneraceae) Ham am elidaceae Hernandiaceae* H ydnoraceae H ydrocharitaceae H ydrophyltaceae H ydrostachyaceae H ypericaceae H ypoxidaceae Icacinaceae lllecebraceae Iridaceae Juncaceae iuncaginaceae Lam iaceae Lauraceae L ecythidaceae L em naceae Lentibulariaceae Liliaceae Linaceae Loasaceae Lobeliaceae Loganiaceae L oranthaceae (s.s.) L oranthaceae (s.l.) (see Viscaceae) L ythraceae Malphigiaceae Malvaceae M artyniaceae* M elastom ataceae Meliaceae M elianthaceae M enispermaceae M esem bryanthem aceae M ontiniaceae Moraceae M oringaceae M usaceae (s. s.) M usaceae (s.1.) (see Strelitziaceae) M yoporaceae* M yricaceae M yrotham naceae Myrsinaceae M yrtaceae N ajadaceae N yctaginaceae N ym phaeaceae O chnaccae O lacaceae Oleaceae Oliniaceae O nagraceae Opiliaceae O rchidaceae O robanchaceae* O xatidaceae Papaveraceae G enera 1 11 5 2 1 1 I - Species & infrasp. taxa 1 90 279 52 2 3 6 3 1 76 267 44 2 3 3 Species G enera Species 5 12 3 1 65 265 47 2 3 4 1 12 4 5 1 7 5 2 1 1 1 1 4 2 3 1 62 271 43 2 3 3 1 12 3 4 t 1 1 4 2 1 4 1 3 11 2 1 5 3 5 43 3 1 37 5 1 6 2 60 2 1 5 5 91 6 8 879 38 2 256 12 1 9 16 949 6 1 141 22 1 1 1 4 1 1 1 6 3 4 44 3 1 31 4 1 6 2 54 2 1 . 5 3 1 3 11 2 1 7 83 6 9 846 19 2 225 11 1 9 18 907 6 1 . 24 . 54 - 12 - - G enera 1 10 5 2 1 1 1 1 4 2 3 - 4 G o ld b latt (1978) D yer (1 9 7 5 ,7 6 )a Indigen. species 1 1» 1 4 1+1* 1 6 3 3+2* 46+ 1* 3 1 3 2 -« « 4+1 * 1 4 2 57+2* 2 1 6 5 11 12 3 I 3 11 3 1 3 0 3 11 2 1 86 6 13 1024 23 2 257 13 1 10 20 1 142 5 1 181 24 42 61 • 80 6 7 858 20 2 225 12 1 10 19 1 066 5 1 141 24 40 58 6+1* 3 15 -* * 2* 4 7+2* 2 7+1* 123 1 4 1 1 2 49 17 161 2 11 17 13 15 2684 1 28 1 1 6 1* 1 1 4 4+3* 1 4+1* 1 2 2 5 1 2+2. 1 54 1» 2 1+2. 2 9 1 8 29 3 16 4 15 4 27 4 34 1 480 2 243 7 - • - - 2 48 12 43 13 139 0 9 14 13 13 2408 1 28 1 1 6 7 3 21 ] 4 9 2 7 122 1 4 1 1 2 27 12 116 0 9 15 12 14 2 304 1 27 1 1 6 7 3 14 4 6 2 7 0 9 I 8 24 3 10 4 14 2 22 4 12 1 439 0 195 1 1 1 1 4 6 1 5 1 2 2 5 1 4 1 54 1 2 2 0 9 1 7 21 3 11 4 11 3 22 3 11 1 439 1 ?200 1 35 13 122 D ifference SL (+14) D (+ 11) D (+21) D (+ 18) D (+31) SL (+ 159) SL (+16) SL (+23) - 9 12 14 13 . - 1 4 1 1 2 - 1 27 1 1 6 . 1 1 4 4 1 4 1 2 2 5 1 3 1 54 9 1 7 8 3 10 4 13 3 22 4 11 1 461 2 5 203 7 . S L (+16) G (+ 22) 627 B othalia 15, 3 & 4 (1985) Fam ily Passifloraceae Pedaliaceae Penaeaceae Peripiocaceae Phytolacca ceae Piperaceae Pittosporaceae Plantaginaceae Plum baginaceae Poaceae Podostem aceae Polygalaceae Polygonaceae P ontederiaceae P ortulacaceae P otam ogetonaceae P rim ula ceae P roteaceae Ptaeroxylaceae Punicaceae* Rafflesiaceae R anunculaccae Reseda ceae R estionaceae R etziaceae R ham naceae R hizophoraceae R oridulaceae Rosaceae R ubiaceae R uppiaceae R utaceae Salicaceac Salvadoraceae Santalaceae Sapindaceae Sapotaceae Sctophulariaceae Selaginaceae Sim aroubaceae Solanaceae Sphenocleaceae Sterculiaceae Stilbaceae Strelitziaceae Tam aricaceae Tecophiliaceae T hym elaeaceae Tiliaceae T rapaceae Trim eniaceae (also M onim iaceae) T urneraceae T yphaceae Ulm aceae U rticaceae V ahliaceae V alerianaceae V elloziaceae V erbenaceae V iola ceae Viscaceae V itaceae X yridaceae ZannicheUiaceae Zingiberaceae Zosteraceae Zygophyllaceae TO TA L (seed plan ts only) List o f Species (1984) Indigen. G enera Species & infrasp. species taxa 4+1» 8 7 8 2+1» 2 1 1 3 171+28» 4 4 4+4* 3+1» 5 1 3 13+1» I 1* 1 7 l+ l* 12 1 8 4 13+4* 59+1» 1 20+1* 1+1» 2 6 13 7 53-«5* 10 1 4+5* 1 7 5 1 1 2 9 4 1 1 4 1 3 11+1« 1 2 2 10+3* 3 1 5 1 5 2 I 8+1* 2076 + 226» 28 43 26 23 7 5 3 11 27 955 5 210 49 5 65 7 8 392 1 1 2 30 5 301 1 203 8 180 236 2 306 10 4 178 29 14 568 218 3 98 1 186 12 5 3 10 199 61 2 1 11 2 5 28 7 2 9 78 10 19 55 7 5 4 1 55 22647 18 34 21 21 5 5 3 4 24 783 5 205 35 3 64 7 8 366 1 0 2 24 3 282 1 159 8 150 207 2 291 6 3 176 29 14 543 214 3 68 1 175 12 5 2 8 189 56 1 1 10 2 5 24 2 2 9 60 9 18 52 7 5 3 1 53 19776 + 204# D yer (1 9 7 5 ,76)a G enera Species 4 8 7 7 3 2 1 1 3 192 4 4 6 3 5 1 3 14 1 1 7 2 12 1 8 4 1 16 58 1 21 2 2 6 13 7 53 9 1 6 1 6 5 1 1 2 9 4 1 1 4 1 3 12 I 2 2 13 3 5 1 4 1 1 7 1853 16 30 21 25 4 6 1 10 23 840 4 207 32 3 62 7 8 365 1 . 3 25 2 193 1 173 8 2 145 232 2 270 5 3 163 25 13 495 206 3 50 1 229 10 5 1 10 165 48 1 1 8 1 5 26 1 2 7 67 8 49 10 5 2 1 62 18369 G oldblatt (1978) G enera Species D ifference 6 13 7 51 10 1 3 1 6 5 1 2 8 4 1 4 1 3 11 1 2 2 9 3 5 1 4 1 1 7 17 29 21 5 6 1 10 21 743 4 201 31 3 56 7 7 336 1 3 25 3 316 1 156 8 2 133 234 2 271 5 3 144 23 14 515 210 3 59 1 169 13 1 10 179 49 1 9 1 5 27 2 2 9 55 8 48 10 5 2 1 46 1859 18327 4 8 7 • 2 2 1 1 3 169 4 4 4 3 5 1 3 14 1 1 7 1 12 1 9 4 1 9 59 1 23 1 2 D & G (+6) D (■♦57) S L (+ 8 ) SL (+30) G (+ 34) D (+14) S L (+17) G (+ 27) SL (+21) SL (+ 32) SL (+48) SL (+ 18) D (+ 54) S L (+24) S L (+8) D (+7) D(-*9) * D yer (1 9 7 5 , 1976) o m itte d species c o u n ts fo r a num ber o f genera. T hese w ere estim ated fro m PRECIS lists d a tin g from th o se years. * N aturalized genera * U ndescribed species B othalia 15, 3 & 4 (1985) 628 species in F abaceae, an increase o f ab o u t 400 species in this family, C O N C L U S IO N S T h e num ber o f species o f seed plants estim ated for th e southern African flora has increased by roughly 2 300 in th e past fourteen years. T he question arises w h ether the present to tal, based o n the List o f Species o f Southern African Plants (1984) will be just as quickly o u td ated . T h ere are th ree sources of changes in the num bers o f recognized taxa: lumping and splitting o f presently recognized taxa, which can eith er increase o r decrease the total; ‘finding’ taxa in the H erbarium as a result o f correctly identifying existing specim ens as undescribed taxa o r as records o f taxa not previously known from southern Africa, which will increase the total; and collecting new o r new ly-recorded taxa in the field, which will also in crease the total. T h e present trend in taxonom y is generally tow ard reduction in th e num ber o f species recognized (B ren an, 1978). C ertain fam ilies, particularly M esem bryanthem aceae, are likely to have th eir num bers o f taxa greatly reduced w hen they are critically revised. T h ere could be a reduction o f over 1 000 taxa in M e sem bryanthem aceae alone. H ow ever, the cases of Asparagus and F abaceae show th a t although there may be a general trend tow ard reduction, and great reductions in som e fam ilies, not every group will have its num ber o f taxa reduced w hen it is revised. T he num bers o f unpublished species and taxa given in T ables 1 and 2 are only those which are so well know n that they have a 'm anuscript nam e' awaiting publication, and they account fo r only 1,4% o f th e to tal num ber o f known taxa. O bser vation o f th e P R E herbarium shows th at th e re a re a great m any specim ens in ‘sp p .’ folders which await critical work by experts, and m any will probably prove to be new o r new ly-recorded taxa. D oubtless th e sam e situation exists in o th er herbaria w ith large holdings o f southern A frican plants. Even if only o n e genus in ten contains a new species, th ere could be a fu rth er 250 species now represented by H erbar ium specim ens. O verall, southern A frica is reasonably well cov ered by plant collections. P R E C IS records show that every whole degree square has som e specim ens re corded, and since the P R E C IS records represent only about 16% o f th e total o f H erbarium specim ens in southern A frica, th e true coverage is probably b e tte r th an the sam ple shown by P R E C IS (G ibbs Russell, R etief & Sm ook, 1984). N evertheless, the arid central and w estern parts o f southern A frica re quire m uch greater collecting efforts in o rd e r for their plants to be as well represented in herbaria as those o f the m ore mesic south and east. B ecause o f th e low er collecting intensity in the dry areas, it is likely th at th ere are m ore new taxa aw aiting discov ery th ere th an in the better-know n mesic areas. For these reasons, changes in the num bers o f taxa known for th e southern A frican flora are bound to occur as a result o f the basic taxonom ic activities o f plant collecting, herbarium curatioo and revision. In th e fu tu re, these changes can be easily m onitored and recorded by updating the List o f Species in P R E C IS, so th at com plete o r partial lists can be printed by com puter, and to tal num bers determ ined. T he total now recognized is so large th at even a change o f 1 000 taxa represents only 4% o f the total flora. T he com parisons m ade here thus may change in detail, but the overall perspective is unlikely to alter. T h e com parative picture outlined here em phasizes the richness o f the southern African flora in term s of high species/area ratios, many vegetation types and high levels o f endem ism , especially in som e o f the largest families and genera. T his richness is unequal led anyw here else in the w orld on a subcontinental scale, and dem ands th a t a high priority be given to the system atic study necessary to understand it. ACKNOW LEDGEM ENTS Mrs W, Roux has patiently checked and proofread all taxon counts from the List o f Species. D r O . A. Leistner, D r P . G oldblatt and M r E . G . H. O liver have m ade valuable criticisms o f th e m anuscript. U lT T R E K SE L Die flora van Suider-Afrika is vir die eerste keer tot by spesies vlak in die List o f Species o f Southern African Plants (Gibbs Russell et al., 1984) ondersoek. Die aantal taksa wat vir Suider-Afrika opgeteken is word vergelyk met dié van taksa wat vir ander dele van Afrika aangegee is, en die grootste families in elke streek word gelys en vergelyk. Die rykdom van spesies in Suider-Afrika word m et dié van ander dele van die wêreld vergelyk. Die aantal genera, spe sies en subspesies vir elke familie in die Suider-Afrikaanse flora word gegee net met vorige tellings deur Dyer (1975, 1976) en Goldblatt (1978) vergelyk. REFEREN CES A D A M S O N . R . S ., 1938. The vegetation o f South Africa. L on d o n : W hitefriars Press. A D A M S O N , R . S, & S A L T E R . T , M ., 1950. The flowering plants o f the Cape Peninsula. C ap e T ow n: Juta. A N D R E W S . F, W „ 1950. The flowering plants o f the AngloEgyptian Sudan. Vol. 1, A rbroath: Buncle. A N D R E W S , F . W ., 1952. The flowering plants o f the AngloEgyptian Sudan, Vol. 2. A rb ro a th : Buncle. A N D R E W S , F. W ., 1956. The flowering plants o f the AngloEgyptian Sudan, Vol. 3. A rb ro a th : Buncle. B R E N A N , J . P. M ,, 1978. Som e aspects o f the phytogeography o f tropica! A frica. Ann. Mo. bot. Gdn 65: 437-478. C L A Y T O N , W . D . & R E N V O IZ E . S .A ., 1982. Poaceae. In R. M , Polhill, Flora o f Tropical East Africa. 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