A revised cytotaxonomy of the genus Tulbaghia (Alliaceae)

CARYOLOGIA Vol. 53, no. 2: 83-112, 2000 

A revised cytotaxonomy of the genus Tulbaghia 

(Alliaceae) 

CANIO G. VOSA* 

Linacre College - Oxford, England. 

In memoriam Dr. L. Codd, eminent South African Botanist and a very dear friend. 

INTRODUCTION 

Because of its validity, especially regarding 

the early history of the genus, it has been decided 

to publish the original introduction by 

VOSA (1975), in a corrected and appropriately 

up-dated version. 

The genus Tulbaghia was created by Linnaeus 

(Mant. Pi. p. 148, 1771) and based on material 

sent to Europe in 1769 by C. Rijk van Tulbagh, 

then Governor of The Cape Province. 

Linnaues, in fact, used the name Tulbagia, 

which was corrected into Tulbaghia by P.D. 

Giseke (Prael. Ord. Nat. PL 274, 1792). The 

first species to be described, and the basis of the 

genus, is Tulbaghia capensis. It is illustrated in 

Jacquin (PI. Hort. Bot. Vindob., 2: 52, pi. 115, 

1772-1773) and, as Tulbaghia alliacea, in the 

Bot. Mag. pi. 806 (1805). The specimen illustrated 

in PL Hort. Vindob. seems to correspond 

* For correspondence: Dipartimento di Scienze Botaniche, 

via Luca Ghini, 5, Pisa, Italy. Fax ++39 050 551345, email: 

caniovosa@tin.it 

Abstract — A revision of Tulbaghia (Alliaceae) has been carried out. The genus includes 

twentyone species and one variety. On the basis of the different karyotypes, the 

genus can be subdivided into seven distinct Groups. Some species have poly-ploid 

forms but no true polyploid species has been found so far. A number of species 

possess heterochromatic segments which may be either mostly distally or 

proximally located. The heterochromatic segments of Tulbaghia are cold sensitive 

and specific fluorochrome staining shows that they are composed mainly of highly 

repetitive sequences of Guanine-Cytidine nucleotides. The possible evolutionary 

role of heterochromatin as well as that of the B-chromosomes, present in some species, 

is briefly discussed. The illustrations include micrographs of the mitotic chromosomes 

as well as colour photographs of the inflorescence of all the species. The typical 

habitat of two of the species is also illustrated in colour. 

Key words: B-chromosomes, constitutive heterochromatin, cytotaxonomy, karyotype 

groups, nucleolar organizers, Tulbaghia. 

to one of the four specimens of Tulbaghia 

present in the Linnaean Herbarium (n° 411.1). 

Two more species of Tulbaghia were described 

ten years later by Linnaeus filius in the Suppl. 

Plant, of 1781 (p. 193-194). The first descrition 

refers to a new species, Tulbaghia attiacea (n° 

411.2 of the Linnaean Herbarium — nectarium 

ori tubi impositum, sexdentatum, obsoletius, obtusum, 

crassus, longitudine limbi). The second 

description refers to a purple flowered plant 

with a three-lobed corona (n° 411.4, Linn. Herb. 

— differt a T. alliacea nectario triphyllo, statura 

minore). It gives the name T. cepacea, but 

includes Linnaeus's T. capensis as a synonym and 

refers also to the PL Hort. Vindob. illustration. 

This synonymy, and the name alliacea given to 

the illustration of T. capensis in the Bot. Mag. (pi. 

806, 1805), together with the annotation by J.E. 

Smith on the T. capensis type specimen (411.1) 

and his note on T. alliacea in Rees Cyclop Roem. 

etSchult. 7: 993 (1819), have been responsible for 

the confusion in the early nomenclature of the 

genus. 

A little known revision of the genus Tulbaghia 

was published by J.E.L. Ave-Lallemant

86 VOSA 

nently coloured. Intermediate types include T. 

coddii and T. natalemis where the perianth segments 

are pigmented but in contrasting colours 

with the corona. Some species such as T. galpinii 

and T. rhodesica represent special cases. T. 

galpinii is a tiny species which resembles a diminutive 

T. capensis with a deeply cut, but not 

very fleshy corona, very thin flaccid leaves and a 

different karyotype which places it in the T. natalensis 

group (Group 5). T. galpinii'is endemic 

of a very restricted area of the Eastern Cape and 

all cultivated collections seem to be self-fertile. 

Tulbaghia rhodesica is a tropical species. It 

has a karyotype similar to that of T. cameroni, 

which comes from the same region, but, in contrast 

with his latter species, has brightly colored 

flowers with broad perianth segments and a very 

short, deeply lobed corona of the same colour. It is 

difficult to define the affinities of T. rhodesica 

which probably represents a very ancient type 

well adapted to tropical light forest conditions. 

Tulbaghia simmleri may represent an extreme 

evolutionary type. It possesses some peculiar 

characters such as its flower morphology 

with somowhat fleshy perianth segments and the 

comparatively small chromosomes (VosA 

1975). Its breeding behaviour in crosses with 

other species is interesting showing rather scarce 

affinity. In fact, although a number of seeds in 

produced in most reciprocal crosses and these 

germinate well, the resulting plantlets seem to 

have a defective root system and fail to grow 

(VOSA 1996c). 

Field observations have shown that population 

structure, in different species, presents some 

interesting features. In optimal conditions, all the 

species with highly coloured or white flower, 

such a T. violacea, T. cominsii, T. natalensis and 

T. coddii, occur often in dense populations 

especially along water courses or in vleis or, as in 

the case of T. simmleri, in light but somewhat 

humid mountain forest. All the species with 

inconspicuosly coloured flowers such as T. 

acutiloba, T. cernua, T. alliacea, T. capensis, etc. 

occur mostly in very small clumps or, more often, 

as single plants at quite a distance from one 

another. This situation may probably reflect a 

kind of strict interaction between flower colour 

and eventual pollinators. 

Taxonomic treatment 

1. Tulbaghia capensis Linnaeus, Mant. 2:148,223 

(1771); Jacq. Hort. Vindob. 2: 52, PL 115 (1772) 

1773);WIUd.,Sp.P1.2:33(1779);Enum.Hort.Berol. 

349 (1809); Baker, J. Linn. Soc. Bot. (London) 11:370 

(1871), Fl. Cap. 6: 406 (1897); Uphof, Herbertia, 10: 

45 (1943); Vosa, Ann. Bot. 34: 66-68 (1975); Burbidge, 

Notes R. hot. Gds. Edinb. 36: 84 (1978). 

TYPE —411.1 (Linn!). 

SYNONYMS — Tulbaghia pulchella Ave-Lall. in Bull. 

Cl. Phys. Math. Acad. Petersb. 3: 204 (1844), nom. 

illegit. 

Tulbaghia alliacea auct. non L. £, Sims, Curtis's 

Bot. Mag. 21: t. 806(1805). 

ICONOGRAPHY — Jacquin in PI. Hort. Bot. Vindob. 2: 

52, PL 115 (1772-1773); Curtis's Bot. Mag. 21: t. 

806, 1805; (VosA 1975, fig. 3, Plate I, fig. 3); Plate 

IA. 

CHROMOSOME NUMBERS — 2n=2x=12; 2n=4x=24; 

2n=6x=36 (karyotype C, group 1, fig. 22b; fig. 1). 

DISTRIBUTION — Endemic to the South Western 

Cape. Diploids and tetraploids are found only in the 

eastern part of the range. 

EXSICCATA — Diploids: CAPE. - 3421 (Riversdale): 

1.5 km E of Heidelberg, Vosa, 294/14 (OXF). 

Figs. 1-21 — Mitotic metaphases in Tulbaghia: in all micrographs, 

the positions of the nucleolar organizing regions are 

indicated by arrows. Fig. 1 — Tulbaghia capensis (2x), karyotype 

C.

CYTOTAXONOMY OF THE GENUS TULBAGHIA 87 

Tetraploids: CAPE. - 3421 (Riversdale): just W of 

Heidelberg, Vosa, 680 (OXF). - 3320 (Montagu) 11 

km W of Drew, Vosa 2459 (OXF). 

Hexaploids: CAPE. - 3318 (Cape Town): Waylands 

Farm, Darling, Vosa 188/2 (OXF); Schwartzvlei 

Farm, Darling, Vosa 189/3, 190/4 (OXF). - 3319 

(Worcester): Karroo Garden on rocky hillside, Vosa 

232/6 (OXF); 8 km S of Villiersdorp, Vosa 143/10 

(OXF). 

Other exsiccata: CAPE. - 3318 (Cape Town): 

Mamre Hills, Malmesbury, Compton 17271 (NBG), 

Baker 1643 (NBG). - 3420 (Bredasdorp), Lewis 

57187 (NBG); Axelson 263, Tinie Versfeld Reserve, 

Darling (NBG); Slangkop, Darling, Rycroft 1778 

(NBG). 

This taxon is the type species of the genus. It is 

confined to the winter rain region of South Africa 

and is distinguished from all others in the genus by its 

deeply lobed and usually very fleshy corona-lobes. 

2. Tulbaghia alliacea L. f. in Suppl. PL: 183 

(1781); Ave-Lallemant, Bull. Cl. Phys. Math. Acad. 

Petersb. 3: 201 (1844); Baker J. Linn. Soc. Bot. London, 

11: 371 (1871), Flora Capensis, 6: 405 (1897); 

Uphof, Herbertia, 10: 48 (1943); Marloth, Fl. PI. S. 

Africa, 4: pi. 27 (1915); Vosa, Ann. Bot. (Rome), 34: 

70-71 (1975); Burbidge, Notes R. bot. Gds. Edinb. 

36: 94-85 (1978). 

TYPE —411.2 (Linn). 

SYNONYMS — Tulbaghia narcissi/lorn Salisb. sensu 

Podr. Stirp. 219 (1796); Tulbaghia inodora Gaertner, 

sensu De Fruct. et Sem. PI. 1: 57 (1833); Tulbaghia 

brachystemma Kunth, sensu Enum. Pi. 4: 483 (1833).. 

ICONOGRAPHY — (VosA 1975, fig. 4, Plate II, fig. 3); 

Plate IB. 

CHROMOSOME NUMBERS — 2n=2x=12; 2n=6x=36 

(karyotype C, fig. 2). 

DISTRIBUTION — This species, one of the oldest 

established in the genus, is found only in a rather restricted 

area of the winter rainfall region of South Africa. 

It is very variable in size. A population with 

2n=6x=36 (hexaploid) is found in the middle of Cape 

Town, in Rondebosch Common. Tulbaghia alliacea is 

very distinct from all other species of the genus in its 

morphology and distribution which partly shares with 

T. capensis. 

EXSICCATA — Diploids: CAPE. - 3318 (Cape 

Town): dry slopes above Camp's Bay, Vosa 61 (OXF); 

moist sandy meadow, among large stones, Botanical 

Reserve, Stellenbosch, Vosa 242/9 (OXF); - 3418 

(Simonstown): Cape Flats, in fine sandy soil, Vow 48 

(OXF). 

Fig. 2 — Tulbaghia alliacea (2x), karyotype C. 

Hexaploids: CAPE. - 3318 (Cape Town): Rondebosch 

Common (Cape Town) in wet peaty soil on 

low lying ground, Vosa 244/11. 

Other exsiccata: CAPE. - 3218 (Clanwilliam): 

slopes along the river Kloof, Olifants Rivier Mountains, 

Esterhuysen 15303 (BOL); - 3318 (Cape Town): 

Rondebosch Common, Leigh ton 1619 (NBG); 

Stellenbosch, Boucher 145 (STE); - 3418 (Simonstown): 

Slangkop, Pond 1497 (BOL); 3419 (Caledon): 

steep bank above rocks, Riviersonderend Mts. 

near Greyton, Esterhuysen 20754 (BOL). 

3. Tulbaghia cernua Ave-Lallemant, Bull. Class 

Phys. Math. Acad. St. Petersb. 3: 202-203 (1844), 

Ind. Sem. h. Petr. 9: 25 (1844), Linnaea, 18: 222 

(1844); Vosa, Ann. Bot. (Rome), 34: 75-77 (1975); 

Vosa, J.S. Afr. Bot. 47: 57-61 (1981). 

TYPE — Ave-Lallemant 73.6 (LE!).

88 VOSA 

SYNONYMS — Tulbaghia alliacea var. affinis Baker, 

sensu Link Enum. alt. 1: 310 (1821); Tul-baghia 

alliacea forma a Drege sensu Kunth Enum. 4: 483 

(1843); Tulbaghia ludwigiana Harv. forma macrior, 

sensu Kunth. Enum. 4: 483 (1843); Tulbaghia 

campanulata N.E. Br. in Kew Bull. 175: 136 (1901); 

type: CAPE. - Queenstown Division: mts. near 

Queenstown, Galpin 1660 (K, holo!). 

ICONOGRAPHY — Batten & Bokelmann in "Wild 

Flowers of the Eastern Cape", p. 10, pi. 6, fig. 7 

(1996); Vosa, 1975, fig. 3, Plate I fig. 3; Vosa in J.S. 

Afr. Bot. 47: 57-61 (1981); Plate 1C. 

CHROMOSOME NUMBERS — 2n=2x=l2; 2n=4x=24 

(karyotype A, fig. 3). 

DISTRIBUTION — Eastern Cape to Northern 

Transvaal. The diploids have been found so far only 

in the southern part of the range. 

EXSICCATA — Diploids: NATAL. - 3030 (Port 

Shepstone): High Flats, Vosa 413/32 (OXF); CAPE. - 

3129 (Port St. John): 3.5 km S of Flagstaff, Vosa 14 

(OXF); - 3130 (Port Edward): between sand dunes, 

Vow 416/33 (OXF). 

Tetraploids: TRANSVAAL. - 3229 (Pieters-burg): 

Haenetsburg, near Old German Mission, Vosa 58 

(OXF). KWAZULU-NATAL. - 2931 (Stan-ger): 

Groenberg (Inanda), Vosa 406/31 (OXF); -2929 

(Underberg): Estcourt, Van der Merwe 2544 (PRE); 

CAPE. - 3326 (Grahamstown): Haye's Railway 

Sidings, Vosa 392/28 (OXF); 1.5 km NE of East 

London, on the side of the road to Gonubie Park, 

Vosa 306/16 (OXF). 

Other exsiccata: CAPE. - 3227 (Stutterheim): 

King Williams Town, Dyer 254 (PRE); - 3425 (Port 

ELizabeth): near the town, Eayliss 1567 (K). 

VERNACULAR NAMES — Southern Sotho: Moe-lela. 

Xhosa: Ivimba 'Mpunzi'. KwaZulu: Si'kwa. 

Tulbaghia cernua is a variable species and the two 

cytological forms overlap especially in the size of the 

flowers and of the leaves. Its main characteristic is 

the urceolate or barrel-shaped fleshy corona, the lanceolate-obtuse 

perianth segments and the large rhizomatous 

base of the bulb. It can be distinguished 

easily from T. acutiloba (with which it shares the 

character of the corona) by the very different leaves 

(very much narrower and tapering in T. acutiloba], 

by the larger corm and by the general shape of the 

flowers which are smaller, narrower and elongated in T. 

acutiloba. 

T. cernua has been usually confused with T. alliacea 

and indeed many herbarium collections bear this 

name. However, it is very distinct and T. alliacea is 

found only in the winter rainfall region of the Cape 

Province. Besides distribution, the differences be- 

tween T. alliacea and T. cernua include important details 

of morphology and of karyotype. An illustration 

agreeing with T. cernua appears under the name of T. 

alliacea in: "Wild Flowers of the Eastern Cape Province" 

by Batten and Bokelmann, p. 10, PI. 67, fig. 7 

(1966). No indication is given about the location of 

the original plant except that the species is known to 

occur throughout the Cape. 

Fig. 3 — Tulbaghia cernua (2x), karyotype A. 

4. Tulbaghia ludwigiana Harv. Bot. Mag. 64: t. 

3547 (1837); Dietr. Synops. 2: 1090 (1840); non 

Kunth Enum. 4: 482 (1843); Ave-Lallemant, Bull. Cl. 

Phys. Math. Acad. Petersb. 3: 203 (1844); Uphof, 

Herbertia, 10: 46 (1943); Schnizlein, Icon. gr. 1: t. 55 


Burbidge, Notes R. bot. Gds. Edinb. 36: 86-88 

(1978). 

TYPE — Harvey in Bot. Mag. 64: t. 3547 (1837): 

iconotype! 

The species has been described and illustrated 

from cultivated material collected by Mr. Zeir (Zeyhr?) 

from the border of "Kafferland" and not known to 

have been preserved as herbarium specimen. The 

beautiful illustration in Bot. Mag. 64 (1837) has been 

selected as iconotype.


Fig. 4 — Tulbaghia ludwigiana (2x), karyotype A. 

ICONOGRAPHY — (Harvey, 1837, Bot. Mag. 64: t. 3547; 

Vosa, 1975, fig. 6, Plate IV, fig. 3); Plate ID. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotype A, 

fig. 4). 

DISTRIBUTION — Eastern Cape Province. 

EXSICCATA — CAPE. - 3327 (Peddie): in grassy 

fields at edges of Euphorbia scrub, south bank near 

the mouth of the Great Fish River, Vosa 398/29 

(OXF), Vosa 2171 (OXF); - 3326 (Grahamstown): 

W of Eraser's Camp, Barker 6993 (NBG). 

This species is closely allied to T. cernua but it is 

easily distinguished by the very broad, lorate leaves 

which have a more or less distichous arrangement 

and lie flat on the ground. 

5. Tulbaghia dtegeana Kunth in Enum. PI. 4: 483 

(1843); Ave-Lall., Bull., Cl. Phys. Math. Acad. Petersb. 

3: 203 (1844); Baker, J. Linn. Soc. Bot. (London) 

11: 371 (1871) and El. Cap. 6: 405 (1897); Uphof, 

Herbertia, 10: 46 (1943); Vosa in Ann. Bot. 

(Rome), 34: 77-79 (1975); Burbidge, Notes R. bot 

Gds. Edinb. 36: 88-89 (1978). 

TYPE — CAPE. - 3018 (Kamiesberg): near Lilyfontein 

and Ezelfontein, Drege 2658 (BM). 

ICONOGRAPHY — (Vosa, 1975, fig. 7, Plate V, fig. 3); 

Plate IE. 


group 2, fig. 22b; fig. 5). 

DISTRIBUTION — Southern and Western Cape 

Province. 

EXSICCATA — CAPE. - 3118 (Vanrhynsdorp): 

bottom of Vanrhysdorp Pass, Vosa 62 (OXF); - 3319 

(Worcester): Lemoenspoort, Vosa 1971 (OXF); - 

3320 (Towsrivier) Hardy 4228 (NBG); - 2917 

(Springbook): Spektakel, Barker 1912 (NBG); 

Springbook, Barker 6443 (NBG);-3017 (Hondeklipbaai): 

Komaggas Mts., Hall 900 (NBG); - 3119 (Calvinia): 

Botterkloof, Compton 9642 (NBG); - 3219 

(Wupperthal): near Wupperthal, Hall s.n. (NBG); 

Cap. B. Spei, Drege 1575 (K). 

A very distinct species found around Wupperthal 

and Worcester northwards to Springbook. It is 

easily distinguished from all other species by the very 

short and fleshy corona and by the perianth segments 

all inserted at the same level at the base of the 

corona. 

Fig. 5 — Tulbaghia dregeana (2x), karyotype A.

90 VOSA 

6. Tulbaghia tenuior Krause & Dinter, Bot Jahrb. 

45: 1411 (1910); Uphof, Herbertia, 10: 50 (1943); 

Vosa, Ann. Bot. (Rome) 34: 80-82 (1975); 

Burbidge, Notes R. bot. Gds: Edinb. 36: 91-92 

(1978). 

TYPE — NABIBIA. - 2018 (Grootfontein): near 

Grootfontein, Dinter 790 (MI). 

SYNONYMS — Tulbaghia karasbergensis Glover, 

Ann. Bol. Herb. 1: 104 (1915), type: H.H.W. Pearson 

8192 (BOLD. 

ICONOGRAPHY — (Vosa, 1975, fig. 8, Plate VI, fig. 3); 

Plate IF. 


fig. 6). 

DISTRIBUTION — North Western Cape Province and 

Nabibia. 

EXSICCATA — NABIBIA. - 1918 (Grootfontein): in 

dry vlei at Amrib, Schoenfelder S454, 955 (PRE); 

2120 (Rietfontein): Rietfontein, Vosa 136 (OXF); 

2217 (Windhoek): Windhoek, W. Giess 1598 (PRE). 

CAPE. - 2919 (Pofadder): 3.5 km NE of Pofadder, 

Acocks 21797 (PRE); 2911 (Prieska): Prieska, Bryant 

576 (K). 

Tulbaghia tenuior Krause & Dinter is a very distinct 

species proper of the dry and rocky areas of the 

NW Cape and of Nabibia. It is distinguished by its 

starry flowers with very long and pointed perianth 

segments and by its short, fleshy corona. It is more or 

less deciduous and not very long lived in cultivation. 

In the wild, it flowers between December and 

March. Fig. 6 — Tulbaghia tenuior (2x), karyotype A. 

Fig. 7 — Tulbaghia nutans (2x), karyotype A. 

7. Tulbaghia nutans Vosa, Ann. Bot. (Rome), 34: 

82-84 (1975). 

TYPE — TRANSVAAL. - 2330 (Tzaneen): 

Haenetzburg, C. Thompson Farm, in dry sandy field 

recently burnt, flowering 8/10/1969, Vosa 536/55 

(OXF); Isotype G. 

ICONOGRAPHY — (Vosa, 1975, fig. 9, Plate VII, fig. 

3); Plate IIA. 

7). 

CHROMOSOME NUMBERS — 2n=2x, 4x (karyotype A, fig. 

DISTRIBUTION — Eastern Cape Province and 

Transvaal. 

EXSICCATA — TRANSVAAL. - 2330 (Tzaneen): 

Haenetzburg, in grassy field by the road to Dindinnie, 

Vosa 538/60 (OXF). CAPE. - 3326 (Grahamstown): 

in Euphorbia scrub in sandy soil, between 

Cradock and Vaalkrause Track, Vosa 1536 (OXF); 

Hounslow Farm, on the road to Adelaide, Vosa


2188; Fishriversrand at the crossroad to Adelaide, 

Vow 2197. 

The original description of this species relates to a 

small population found in the Northern Transvaal. 

Further collections near the type locality, and in the 

Kruger National Park, have established the existence 

of small populations and of scattered plants in which a 

tetraploid form predominates. The species is found 

also in the Eastern Cape but here the plants seem to 

be exclusively diploid. Thus, the ploidy situation and 

the geographical range of Tulbaghia nutans appear to 

be similar to that of the more widespread Tulbaghia 

cernua Ave-Lall. which, in the same way, has diploid 

and tetraploid forms and is found from the Eastern 

Cape to the Transvaal. 

8. Tulbaghia macrocarpa Vosa, Ann. Bot. (Rome), 

34: 84-87 (1975). 

TYPE — ZIMBABWE. - 1731 (Harare): amongst 

rocks, in an underdeveloped part of the Botanic Garden, 

Harare, Vosa 132 (OXF), isotype G. 

ICONOGRAPHY — (Vosa, 1975, fig. 10, Plate VII, fig. 

3); Plate HE. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotypeAA, 

fig. 8). 

DISTRIBUTION — Zimbabwe. 

EXSICCATA — ZIMBABWE. - 1731 (Harare): in the 

Botanic Garden, Harare, H.M. Biegel2859. 

Fig. 8 — Tulbaghia macrocarpa (2x), karyotype AA. 

Fig. 9 — Tulbaghia transvaalensis (2x), karyotype AA. 

This species represents one of the Tulbaghia 

types which are usually labelled "alliacea" in most 

herbarium collection. However, it is very distinct 

and, besides its northern tropical distribution, it 

presents several discriminating characters such as the 

very large ovary, which develops in a long capsule 

(hence the specific epithet), and the rather fleshy, 

laterally expanded corona. 

9. Tulbaghia transvaalensis Vosa in Ann. Bot. 

(Rome), 35: 87-90 (1975). 

TYPE — TRANSVAAE. - 2330 (Tzaneen): on 

dry, stony hillside, Gordon McNeil Farm, Unkumani, 

Dindinnie (Ofcolaco), Vosa 486/49 (OXF), 

isotype G.

92 VOSA 

ICONOGRAPHY — (Vosa, 1975, fig. 11, Plate IX, fig. 

3); Plate IIC. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotype AA, 

fig. 9). ,- 

DISTRIBUTION — North Eastern Transvaal. 


Unkumani, G. Mac Neil Farm, Dindinnie, Vosa 4867 

29 (OXF); grassy slopes S of Unkumani, Vosa 1644; 

about 2 km SW of Unkumani, in recently burnt field, 

Vosa 1645 (OXF). 

Tulbaghia transvaalensis is a very distinct species 

found so far only in hilly country in the N.E. Transvaal. 

For a long time it has been confused with T. alliacea 

from which it differs very much in its morphology 

and distribution. Its karyotype and certain morphological 

characters place T. transvaalensis in the 

group of T. macrocarpa Vosa. 

Fig. 10 — Tulbaghia acutiloba (2x), karyotype AB. 

10. Tulbaghia acutiloba Harv., Thes. Cap. 2:51, t. 

180 (1854); Baker, J. Linn. Soc. Bot. (London), 11: 

371 (1871); Baker, Fl. Cap. 6: 404 (1897); Phillips, 

Ann. S. Afr. Museum 16: 300 (1917); Uphof in Herbertia, 

10:46 (1943); De Wild., Sel. Hort. Thes. 3:120 


Burbidge, Notes R. bot. Gds. Edinb. 36: 96 (1978). 

SYNTYPES — EASTERN CAPE. - 3126 (Queestown): 

Cooper 463 (K); 3325 (Somerset East): Boschberg, 

Macowan 1582 (K). 

ICONOGRAPHY — (Vosa, 1975, Plate X, fig. 3); Plate 

IID. 

CHROMOSOME NUMBERS — 2n=2x=l2 (karyotype AB, 

fig. 22a, group 4; fig. 10). 

DISTRIBUTION — Eastern Cape Province northward 

to the Transvaal. 

EXSICCATA — TRANSVAAL. - 2528 (Pretoria): in 

recently burnt vlei, Delmas Road, 6 km S of Pretoria, 

Vosa 431/36 (OXF). NATAL. - 2930 (Pietermaritzburg): 

Chelmsford Park, Vosa 347/21 (OXF); 

2931 (Stanger): Groenberg (Inanda), in short grass at 

the top of the hill, Vosa 405/30 (OXF). CAPE. - 

3126 (Queenstown), by the Dam, in short grass 

among rocks, Vosa 373/24 (OXF). 

A very small, inconspicuous species which flowers 

soon after the first spring rain, often in burnt fields. 

It is found, usually, as widely scattered single plants. 

VERNACULAR NAMES — Southern Sotho: Motsuntsunyane. 

11. Tulbaghia simmleri Beauv., Bull. Herb. Boissier, 

8:988(1908). 

TYPE — Shilouvane, (Transvaal), legit H.A. Junod, 

ex hortus G; specimen and drawings on two 

sheets prepared by G. Beauverd (G, holo!). 

SYNONYMS — Tulbaghia pulchella Barnes, S. Afr. 

Card. 20: 185 (1930), non Ave-Lall.; Tulbaghia daviesii, 

C.H. Grey in Hardy Bulbs, 2: 572 (1938), nom. 

nov. for Tulbaghia pulchella Barnes; Tulbaghia fragrans 

Verdoorn in Fl. PL S. Afr. 11: t. 438 (1931); 

R.A. Dyer in Herbertia, 6: 8 (1939); Uphof, Herbertia, 

10 (1943). Type: E. Transvaal, Pilgrim's Rest 

(2430 - DB) Dientje Farm, just N. of Bourke's Luck 

Pot-holes, Celliers, 8894 (PRE, holo!); Vosa, Ann. 

Bot. (Rome) 34: 92-94 (1975); Vosa, J.S. Afr. Bot., 

46: 109-114 (1980). 

ICONOGRAPHY — Beauverd, 1908 - Bull. Herb. 

Boissier, 8: 988, fig. 2; Barnes (l.c.) fig. A (as T. 

pulchella); C.H. Grey (l.c.) as T. daviesii; Verdoorn 

(l.c.) fig. 438 (as T.fragrans); C. Letty in "Wild Flowers 

of the Transvaal", pl. 10, fig. 3 (1962) as T. fra-


grans; Vosa, 1975 - Ann. Bot. (Rome), 34: 91, pi. XI, 

fig. 3, p. 93, fig. 13 (1975), as T.fragrans; Plate HE. 

CHROMOSOME NUMBERS — 2n=2x=l2 (karyotype A, 

fig. 11). 


Letaba, Codd 9455 (PRE); - 2430 (Pilgrim's Rest): 

The Berg, Vosa 466/43 (OXF); Graskop, Vosa 4687 

45 (OXF); Ofcolaco, Cyprus Farm in light moist for est, 

Vosa 530/53 (OXF); near Ohrigstad, among rocks 

by a stream, Vosa 530/53 (OXF); Bourke's Luck 

Potholes, Graskop, Vosa 542/56 (OXF). 

The misinterpretation of the drawings by G. 

Beauverd (attached to the type specimen of T. 

simmleri (see VOSA 1980), by BURBIDGE (1978), has been 

the origin of the assigning of this epithet to T. 

violacea Harv. 

Tulbaghia simmleri is a very decorative and fairly 

hardy species widely cultivated, usually under the 

name of T. fragrans or T. pulchella, in parks and gardens 

in tropical and sub-tropical countries and in cool 

greenhouses in temperate regions. It is endemic of a 

rather small area of the Transvaal, in the northern 

Drakensberg. It is found as isolated plants on rocky 

ledges and in fairly thick colonies in light 

Fig. 11 — Tulbaghia simmleri (2x), karyotype A. 

Fig. 12 — Tulbaghia galpinii (2x), karyotype D. 

tropical forest, together with Clivia miniata Regel 

and epiphytic orchids (mostly Polystachia sp.) and 

ferns, at moderate altitude. 

12. Tulbaghia galpinii Schl, J. Bot. 34: 383 

(1894); Uphof, Herbertia, 10: 47 (1943); Vosa, Ann. 

Bot. (Rome), 34; 94-96 (1975); Burbidge, Notes R. 

bot. Gds. Edinb. 36: 98-100 (1978). 

TYPE — CAPE. - 3126 (Queestown): Andriesberg, 

Golpin 2179 (PRE, isotype!). 

ICONOGRAPHY — (Vosa, 1975, fig. 14, Plate XII, fig. 

3); Plate IIP. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotype D, 

fig. 12). 

DISTRIBUTION — Eastern Cape. 

EXSICCATA — CAPE. - 3225 (Somerset East): near 

Cradock, Vosa 60 (OXF); Bergquagga Park, Vosa 

66; 34 km S of Somerset East, Vosa 67 (OXF); 48 km 

N of Cradock, barker 7065 (NBG); Bergquagga 

Park, on Bankberg, Liebenberg 7127 (PRE); 43 km S 

of Somerset East, Acocks 16741 (PRE). 

A very small species resembling a diminutive T. 

acutiloba in habit but with a scarcely fleshy, deeply 

lobed corona and known only from a fairly small area 

of the Eastern Cape. It inhabits dry, rocky country in 

grassland or in light scrub often associated with 

dwarf euphorbias. 

The three collections cultivated in Oxford (from 

the Cradock Road, about 11 km from Grahamstown, 

the Bergquagga Park and from near Somerset East) 

are all self-fertile but, under cultivation, usually very 

short lived.

94 VOSA 

13. Tulbaghia leucantha Baker, Flora Capensis, 

6:404 (1897); Uphof, Herbertia, 10: 47 (1943); Vosa, 

Ann. Bot. (Rome), 34: 96-99) (1975). 

SYNTYPES — NATAL. - Near the river Tugela, 

Medley Wood 4408 (K!). CAPE. - Zuurberg (Griqualand 

East), MacOwen 1208) (K!). 

SYNONYMS — Tulbaghia dieterlenii Phillips, Ann. S. 

Afr. Museum 16: 300 (1917), type: Leribe (Lesotho), 

Dieterlen 361 (BM). 

Tulbaghia friesii Suesseng, Trans. Rhodes Sci 

Ass. 43: 76 (1957). 

Tulbaghia rhodesica Weimark, Bot. Not. 90: 169 

(1937) nom. illeg. 

ICONOGRAPHY — (Vosa, 1975, fig. 15, Plate XIII, fig. 

3); Plate IIIB. 

CHROMOSOME NUMBERS — 2n=2x-12; 2n=4x=24 

(karyotype D, fig. 13). 

DISTRIBUTION — The diploid forms occur from the 

NE Cape, Lesotho to the N Transvaal while the 

tetraploids occur from N Transvaal to Swaziland and 

Zimbabwe. In the Transvaal, from around Warmbad 

and Pietersburg and the Kruger Park, there seems to 

be an overlap between the two forms. 

EXSICCATA — Diploids: TRANSVAAL. - 2430 

(Pilgrim's Rest): Blyde's River Canyon at 'One Mile 

Walk', in short turf among large rocks, Vosa 54/1 

(OXF); - 2528 (Pretoria): among rocks near Warmbad, 

De Wet 3799 (PRE); in acid, peaty vlei near the 

roadside at Delmas, Vosa 430/35 (OXF); Between 

Kestell and Bethlehem by the Eland's Bridge, Vosa 

1704 (OXF); on rocks in shallow soil by the road 

from Maseru to Ladibrand, Vosa 1711 (OXF). 

Tetraploids: ZIMBABWE. - 1731 (Harare): 

Groombdrige, H.M. Eiegel 2860 (SRGH); TRANS- 

VAAL. - 2431 (Acornhoek): in dry sandy soil in open 

woodland, 10 km N of Skukuza, Vosa 449/38 

(OXF). 

Other exsiccata: TRANSVAAL. - 2528 (Pretoria): 

Delmas Road, Codd 2182 (PRE); 10 km W of 

Hammanskraal, Leach 10659 (PRE). SWAZILAND. - 

2531 (Mbabane): near Bulunga Poort, Manzini, 

Compton 31754 (NBG). ORANGE FREE STATE. - 

2828 (Bethlehem): Hoogland, Liebenberg 7291 

(PRE). CAPE. - 3028 (Matatiele): Mt. Fletcher, Killick 

& Marais 2076 (PRE). NATAL. - 3029 (Zuurberg): 

MacOwen & Bolus (1208) (K); Medley Wood 

1200 (K). 

VERNACULAR NAMES — Southern Sotho: leta la phofu. 

Tulbaghia leucantha is found widespread from 

the N E Cape to N Transvaal, Swaziland and Zimba- 

bwe. It is very variable even within the same population 

in the size of the flowers, in the colour of the corona 

and in the leaves which can vary from very this 

and grass-like to 5-8 mm broad. Overall, the proportion 

of thin-leaved plants, in a given population, seems to 

increase with altitude. The tetraploid forms are much 

less variable than the diploids but on the whole they 

have slightly larger flowers. In any case, most of the 

other characters in both forms, overlap in such a way 

that it is not possible to distinguish them with certainty 

in the field, or in herbarium specimens. 

The morphological variability of Tulbaghia leucantha 

over its considerable range, has resulted in the description 

of new species, mainly based on herbarium 

specimens, by various authors. However, all the forms of 

the species have a very uniform karyotype and are 

interfertile. In any case, the epithet T. leucantha, used 

by Baker in 1896, has priority over T. dieterlenii Philips 

(1917) and over T. friesii Suesseng (1957). 

The name T. rhodesica Weimark (1937), cited as a 

synonym of T. leucantha, is illegitimate because it has 

been already used for a very different species (T. 

rhodesica R.E. Fries, 1916). 

Fig. 13 — Tulbaghia leucantha (2x), karyotype D. Note the 

small satellite intercalary to one of the nucleolar organizer 

thread (double arrow).


14. Tulbaghia natalensis Baker, Card. Chron. 3, 

9: 668 (1891); Baker, Fl. Cap. 6: 405 (1897), Natal 

Plants, 1: t. 29 (1899); Uphof, Herbertia, 10: 49 

(1943), Fl. PI. S. Africa, 25: 979 (1945); Vosa, Ann. 

Bot. 34: 99-101 (1975); Burbidge, Notes R. bot. Gds. 

Edinb. 36: 89 (1978). 

TYPE — NATAL. - 2529 (Underberg): in swamps, 

near Moorivier, Medley-Wood 4045 (NH!). 

ICONOGRAPHY — (Vosa, 1975, fig. 16, Plate XIV, 

fig.3);PlateIIIA. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotypeD, 

fig. 14). 

DISTRIBUTION — KwaZulu-Natal. 

EXSICCATA — NATAL. - 2929 (Underberg): 

Thabamhlope, Estcourt, in acid-peaty pasture, Vosa 

421/34 (OXF); 28 km S of Nottingham Road, Codd 

10582 (PRE); 8 km N W of Nottingham Road, Van 

derMerwe 10581 (PRE); Impendhale, W. Marais 934 

(PRE); Moorivier, Stainbank 3845 (NH); Giant's 

Castle, Edwards 2174 (NU); Thabamhlope, Estcourt, 

Van der Merwe 2585 (PRE). 

A rather attractive species with showy white 

flowers, found in vleis and in marshy habitats. It is 

endemic of Natal and it is almost hardy and easy to 

cultivate in acid compost. 

Fig. 15 — Tulbaghia verdoorniae (2x), karyotype D. 

Fig. 14 — Tulbaghia natalensis (2x), karyotype D. 

15. Tulbaghia verdoorniae Vosa & Burbidge, 

Ann. Bot. (Rome), 34: 101-104 (1975). 

TYPE — TRANSKEL. - 3228 (Willowvale): between 

Mpozolo and Mendu in wet soil, 27-01-1966, 

R. Wood 161, NU 36321 (holo!). 

The name Tulbaghia carnosa, published by R.B. 

BURBIDGE (1978), may be considered a synonym of this 

species. 

ICONOGRAPHY — (Vosa, 1975, fig. 17, Plate XV, fig. 

3); Plate IIIC. 


fig. 15). 

DISTRIBUTION — Eastern Cape, Transkei. 

EXSICCATA — CAPE. - 3228 (Butterworth): in a 

recently burnt vlei, near the railway at Mpuluswa, 15 

km NE of Butterworth (-AA), 07-10-1977, Vosa 

1599 (OXF); on a tussocky, rocky hillside, in pockets 

of wet soil, near the side of the road near Idutywa (- 

AB), 07-10-1977, Vosa 1601 (OXF). 

This species has been cultivated for many years in 

Botanic Gardens of Kew, Oxford and Edinburgh and 

it appears in the respective herbaria as either T. 

capensis or T. alliacea. Vosa, in his two cytological

96 

A 

papers (1966a,b), misapplied the name T. dregeana. 

to this species which has been known under this 

epiteth until 1975. 

Two further collections of Tulbaghia verdoorniae 

have been made in October 1977 by Vosa (1980), 

not very far from the original 1966 collection of R. 

Wood (161, NH 36321), which now represents the 

type. 

The species as been named in honour of the renown 

South African Botanist, Miss I.C. Verdoorn. 

Following the current nomenclatural rules in the 

case of a female person, it has been found proper to 

amend the name verdoornia, as in the original description, 

into verdorniae. 

16. Tulbaghia coddii Vosa & Burbidge, Ann. 

Bot. (Rome), 34: 101-106 (1975). 

TYPE — TRANSVAAL. - 2430 (Pilgrim's Rest): 

Graskop, 1400 m. L.E. Codd 6746 (PRE). 

The name T. poetica, published by R.B. BUR- 

BIDGE (1978), may be considered a synonym of this 

species. 

ICONOGRAPHY — (Vosa, 1975, fig. 18, Plate XVI, fig. 

3); Plate HID. 


fig. 16). 

DISTRIBUTION — Endemic to the Northern 

Transvaal. 

EXSICCATA — TRANSVAAL. - 2430 (Pilgrim's 

Rest): in a small bog just above Lisbon's Falls, Vosa 

496/52 (OXF); very near God's Window, Vosa 5317 

54 (OXF); in short humid turf by the road side 4.5 

km from Graskop towards God's Window, Vosa 

464/41 (OXF); near Lisbon's Falls, Graskop, Brent 

134 (E); marshy ground near Pilgrim's Rest, Davidson 

68 (E). 

A very attractive small species related to Tulbaghia 

natalensis. It has bright green, very thin leaves 

and pink to white flowers with a yellow-green corona 

and a sweet smell reminiscent of that of Narcissus 

tazetta. It grows in bogs or in wet turf in acid conditions, 

sometimes associated with species otDrosera. 

17. Tulbaghia montana Vosa, Ann. Bot. (Rome) 

35:106-109(1975). 

TYPE — NATAL. - 2929 (Underberg): S.E. ridge of 

Giant's Castle, Vosa 70 (OXF), isotype G. 

ICONOGRAPHY — (Vosa, 1975, fig. 19, Plate XVII, fig. 

3); Plate IIIE. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotype 

E, group 6, fig. 22b; fig. 17). • - 

Fig. 16 — Tulbaghia coddii (2x), karyotype D. 

Fig. 17 — Tulbaghia montana (2x), karyotype E. (x 2000). 

VOS


DISTRIBUTION — This small high mountain species is 

apparently confined to the South Eastern slopes of 

the Natal Drakensberg. It is fairly hardy and very 

distinct from the other species of the genus especially 

on account of its dark-green leaves which have 

glaucous undersides, a green midrib and inward 

rolled edges. However, it may be confused with T. 

acutiloba Harv., especially in scarcely annotated and 

badly dried herbarium specimens. 

EXSICCATA — NATAL. - 2929 (Underberg): 

Drakensberg, on the south eastern ridge of Giant's 

Castle, growing in short grass in rather humid black 

soil at the foot of large rocks, Vosa 70 (OXF), Burtt 

5687 (E); without a precise locality but still on the 

south eastern Drakensberg, Buchanan 3899 (E); 3 

km N of Rama's Gate, Bayliss 7831, 7781 (GRA). 

18. Tulbaghia cametoni Baker, Jour. Bot. 16: 321 

(1878); Uphof, Herbertia, 10: 48 (1943); Vosa, Ann. 

Bot. (Rome), 34: 11041 (1975). 

TYPE — Tanzania, shores of Lake Tanganika, 

Cameron 2175 (K!). 

ICONOGRAPHY — (Vosa, 1975, fig. 20, Plate XVIII, 

fig. 3); Plate IIIE. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotypeAF, 

fig. 18). 

EXSICCATA — ZAMBIA. - 1024 (Mwinilunga 

distr.): Richards 16885 (K); 1426 (Mumbwa distr.): 

Mumbwa, Maculay 982 (K); - 1428 (Chilanga distr.): 

Mt. Makulo, Vosa 59 (OXF); - 1626 (Choma distr.): 

near the road from Choma to Itulo, Vosa 68 (OXF). 

Tulbaghia cameroni Bak. is one of the tropical 

species of the genus and is very variable, with some 

plants resembling the tetraploid forms of Tulbaghia 

leucantha. Examination of the various herbarium 

specimens, does not allow a clear view of the characteristics 

of this species and more field work is necessary 

to understand its geographical distribution. 

19. Tulbaghia rhodesica R.E. Fries, Wiss. Ergebn. 

Schwed. Rhodesia-Kongo Exped. (1911/ 1912) 1: 

227-228 (1916), non Tulbaghia rhodesica Weimark, 

Bot. Not. 169 (1937); Vosa, Ann. Bot. (Rome) 34: 

111-113 (1975); Burbidge, Notes R. hot. Gds. 

Edinb.36:97(1978). 

TYPE — Zambia. - 828 (Kawambwa distr.): Kalungwisi 

River, just S of Katwe, R.E. Fries 1911 (LD). 

ICONOGRAPHY — (Fl. PL S. Africa, 35: t. 1383; 

Vosa, 1975, fig. 21, p. 111); Plate IVA. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyotypeAF,fig. 

19). 

Fig. 18 — Tulbaghia cameroni (2x), karyotype AF. Note the telocentric 

B-chromosome (star). 

DISTRIBUTION — Zambia, Southern Tanzania. 

EXSICCATA — ZAMBIA. - 830 (Mporokoso distr.): 

in open bushland in dry sandy soil, near Mporokoso, 

Vosa 74 (OXF); Mporokoso, Richards 13717 (K); 

Chilongwelo, Richards 2196 (K). 

Tulbaghia rhodesica is a very beautiful species, 

found in Zambia and in S. Tanzania. In the colour of 

Fig. 19 — Tulbaghia rhodesica (2x), karyotype AF.

98 VOSA 

its flowers it resembles T. violacea but cytologically it 

belongs to the T. cameroni group. It is a very desirable 

species from the horticultural point of view but it 

is deciduous and has proved to be somewhat difficult 

to cultivate, growing very slowly and flowering 

erratically. 

20. Tulbaghia violacea Harvey in Bot. Mag., 64: t. 

3555 (1837); Vosa, Ann. Bot. (Rome), 34: 113-117 

(1975) as Tulbaghia cepacea L.f.; Vosa, Jl. S. Afr. 

Bot., 46: 109-114(1980). 

TYPE — Described and illustrated from cultivated 

material in the Ludwigsburg Garden, Cape 

Town, by Harvey in Bot. Mag. 64, t. 3555, 1837 

(iconotype!). 

SYNONYMS — T. cepacea L.f. in Suppl. PL 194 

(1781) quoad descr. excl. syn., nom. illeg. type: 411/4 

(LINN); Omentaria cepacea Salisb. Gen. PL Fragm. 

87 (1866). 

ICONOGRAPHY — (Harvey I.e., Batten & Bokel-mann 

in "Wild Flowers of the Eastern Cape", p. 10, pi. 6, 

fig. 6,1966; Vosa, 1975, fig. 22, Plate XX: figs. 3, 4,5,22 

and 23, incl. var. as T. cepacea L.f.); Plate IVB. 

For the legitimacy of the name Tulbaghia violacea 

Harvey, see VOSA (1980). 

CHROMOSOME NUMBERS — 2n=2x=12 (karyo-type A, 

fig. 20). 

DISTRIBUTION — From the Transkei to the SW Cape 

near Port Elizabeth. 

EXSICCATA — CAPE. - 3227 (Stutterheim): Mt. 

Coke, near King William's Town, Compton 144/45 

(NBG); - 3228 (Butterworth): at the mouth of the 

river Kei, Compton 17674 (NBG); - 3224 (Steytlerville): 

in rocky scrubland by the river Gantoos at 

Hankey, Vosa 371/23; 3323 (Willowmore): Zuurberge 

Poort, Barker 7132 (NBG); - 3326 (Grahamstown): 

in bed of dry stream among rocks in the 

Bloukrans, Vosa 354/22,376/26 (OXF); - 3327 (Peddie): 

amongst stones in the bed of the river Buffalo, 

near East London, Vosa 308/17 (OXF); - 3424 (Humansdorp): 

Babeljouw River, Compton 24041 (NBG); 

near Humansdorp, Britten 6513 (GRA). 

VERNACULAR NAMES — KwaZulu: Icinzini. 

All the populations of Tulbaghia violacea, object 

of the present study, show a kind of uninterrupted 

variability and include a number of forms which reasonably 

might be given the rank of varieties. However, 

field experience and careful observations during 

a number of flowering seasons, all over the range of 

the species and also on cultivated plants, have shown 

that considerable differences exist even between 

plants of the same population. Such differences 

include, besides the dimension of the plants, 

Fig. 20 — Tulbaghia violacea (2x), karyotype A. 

also the size of the flowers and the length and shape 

of the lobes of the corona. In a previous account by 

VOSA (1975), one each of the most common forms 

from two fairly large populations, located 200 km 

apart, have been described. A perusal of the descriptions 

shows indeed that the two forms differ, but only 

in some small details. Herbarium specimens, perforce 

limited in number, do not represent the intrinsic 

variability of Tulbaghia violacea populations and 

cannot be taken as the basis for the creation of the 

taxonomic rank of varieties. The exception is the 

establishment of the var. maritima by VOSA (1975). 

This variety is quite distinct morphologically from the 

typical species and exists in fairly small but very 

homogeneous populations with a rather restricted 

habitat near the sea. 

20a. Tulbaghia violacea Harvey var. maritima 

Vosa, Ann. Bot. (Rome) 34: 113-117 (1975). 

TYPE — CAPE. - 3325 (Port Elizabeth): Gantoos 

River Mouth, Vosa, 290/12 (OXF) isotype G. 

ICONOGRAPHY — (Vosa 1975, fig. 23, Plate XX, fig. 

5, Plate IV, fig. 20a, as T. cepacea var. maritima); Plate 

IVC. 

CHROMOSOME NUMBERS — 2n=2x-12 (karyotype A). 

EXSICCATA — CAPE. - 3224 (Humansdorp): near 

the Krom River mouth, Vosa 63 (OXF); - 3323


(Willowmore): Twee River mouth in sandy soil, Vosa 

65 (OXF). 

This variety of T. violacea Harv. has been found, 

so far, only near the sea-shore hence the epithet maritima. 

It differs from the type species by its broader, 

shiny, dark-green leaves without a prominent central 

vein. It has somewhat flat flowers with very small 

coronal lobes. All the collections are very similar to 

one another and self-fertile. 

21. Tulbaghia cominsii Vosa, Jl. S. Afr. Bot. 45: 

127-132 (1979). 

TYPE — CAPE. - 3227 (Stutterheim): in kommit-tee 

soil on rocky ground, often associated with Ner-ine 

filifolia Baker, near Tamaha, 17 km from King 

William's Town (-BD), 28-9-57, D. Comins 1718 

(RU 10586). 

ICONOGRAPHY — (Vosa, 1979, fig. 3, 4); Plate IVD. 

CHROMOSOME NUMBERS — 2n=2x=12 (karyo-typeAA, 

fig. 21). 

DISTRIBUTION — Founds so far only around the type 

locality. 

EXSICCATA — Vosa 1568, 28-10-77 (OXF); Vosa 

1781, 12-11-77 (OXF), on both sides of the road 

from King William's Town near Tamaha. 

This attractive little species is related to T. violacea 

Harv. but differs in its very thin, glaucous leaves, 

white or very rarely pinkish flowers with a pink or 

purplish perianth tube and a different karyotype. The 

glaucous colour of the leaves is very characteristic and 

is due to the stomatal pores, which appears as tiny, 

white streaks under low magnification. 

The chromosome complement of T. cominsii is 

very interesting: with its four nucleolar chromosomes 

of type A, it resembles that of T. transvaalensis 

Vosa and T. macrocarpa Vosa, two taxa quite different 

from the present species whose morphological 

affinities lie with T. violacea. This situation, which 

parallels that of T. rhodesica Fries and T. cameroni 

Bak. (see Vosa, 1975, p. 52), may be an indication of a 

two-track evolution in which the karyotype and the 

phenotype seem to evolve indipendently and, in this 

case, along established pathways. 

The cytology of Tulbaghia 

All the species of Tulbaghia so far studied 

have a fairly uniform anatomy and embryology. 

The chief characters of taxonomic value are the 

morphology of the flowers but vegetative features 

and geographical distribution have their 

importance in the differentiation of the species. 

Fig. 21 — Tulbaghia cominsii (2x), karyotype AA. 

One typical characteristic of the genus is the 

uniformity of the basic chromosome number 

which is x=6 for all the species. A few taxa have 

polyploid forms but not true polyploid species 

has been found as yet. Telocentric B-chromosome 

of various sizes have been found in four 

species (Table 2). 

The chromosome complement of Tulbaghia 

is composed of sub-median chromosomes which 

are among the largest in the plant kingdom: the 

length of the haploid complement vary from 56.5 

to 94.3 μm (Table 1, Figs. 22a and b). 

The karyotypes differ mainly in the type or 

combinations of types of nucleolar chromosomes 

and on this basis the genus may be subdivided 

into seven group of species (Table 1). 

Another character of the chromosome complement 

of some species of Tulbaghia is the presence 

of cold sensitive heterochromatic segments 

(H-segments). Fluorochrome staining has 

stablished that the H-segments of Tulbaghia are 

composed of DNA containing highly repetitive 

sequences of Guanine-Cyti-dine (GC) 

nucleotides (see Figs. 23 and 24).

100 

A 

Fig. 22a — Diagrammatic representation of the six nucleolar 

chromosome types (N.O.R.'s) in Tulbaghia drawn according to 

their relative size. 

Type A: occurs in 13 species and is a medium sized chromosome 

with a submedian centromere and with the N.O.R. located 

in an intercalary position. 

Type B: occurs, in combination with type A, only in one species 

(T. acutiloba Harv.) and is the second longest of the complement: 

the N.O.R. is located in the short arm in an intercalary 

position. 

Type C: occurs in two species (T. capensis L. and T. alliacca L. 

f.) and is a subterminal chromosome and the second longest of 

the karyotype. The N.O.R. is located distally on the short arm. 

Type D: occurs in five species and is usually the shortest of the 

complement with a subterminal centromere. The N.O.R. is located 

terminally on the short arm and often carries an heterochromatic 

"satellite" of various sizes. 

Type E: occurs in one species (T. montana Vosa) and is a medium 

sized chromosome with a submedian centromere. The 

N.O.R. is located terminally on its long arm. Type F: occurs, in 

combination with type A, in two species (T. rhodesica Fries and 

T. cameroni Baker) and is the longest of the complement with a 

submedian centromere: the N.O.R. is located in an intercalary 

position very near the centromere. 

Fig. 22b — Diagrammatic representation of the haploid karyotypes 

of Tulbaghia. One species for each karyotype is illustrated 

and drawn in accordance with its relative size). Group 1: 

T. capensis L. (karyotype C); Group 2: T. dregeana Kunth 

(karyotype A); Group 3: T. transvaalensis Vosa (karyotype AA); 

Group 4: T. acutiloba Harv. (karyotype AB); Group 5: T. coddii 

Vosa & Burbidge (karyotype D); Group 6: T. montana Vosa 

(karyotype E); Group 7: T. rhodesica Fries (karyotype AF). 

VOS

:YTOTAXONOMY OF THE GENUS TULBAGHIA 101 

Plate I. (a) T. capensis L. (x2). (b) T. alliacea L. f. (x2). (c) T. cernua Ave-Lall. (x2). (d) T. ludwigiana Harvey (x2). (e) T. dregeana 

Kunth (x2). (f) T. tenuior Krause & Dinter (x2). 

Description of the karyotypes (Table 1) 

Group A (six species) - The karyotype consists 

of five submedian chromosomes one of 

which carries the nucleolar attachment in a 

proximal position on the long arm and one almost 

subterminal chromosome which is the second 

largest of the complement. In a very few 

plants in certain populations, both in the diploid 

form of T. cernua and in T. ludwigiana, the short 

arm of this latter chromosome varies in length, 

sometimes reaching the size of the shorter arm 

of the second largest chromosome of the Group 

D karyotype. However, in all such cases the 

additional chromosome segment is completely 

heterochromatic.

102 VOSA 

Group AA (three species) - The karyotype is 

overall similar to that of group A but there are 

two nucleolar chromosomes of type A. 

Group AB (one species) - The karyotype 

consists of six submedian chromosomes. The 

second largest carries the nucleolar attachment in 

a proximal position on its short arm (nucleolar 

chromosome type B). There is another nucleolar 

chromosome of type A. 

Group AF (two species) - The karyotype is 

overall similar to that of group A, but the long est 

of the complement carries a nucleolar attachment 

in a very proximal position on its long arm 

(nucleolar chromosome type F). There is another 

nucleolar chromosome of type A. 

Group C (two species) - The karyotype resembles 

that of the species of group A, but the 

nucleolar attachment is carried distally on the 

short arm of the second largest chromosome. 

Plate II. (a) T. nutans Vosa (x3). (b) T. macrocarpa Vosa (x2). (c) T. transvaalensis Vosa (x2). (d) T. acutiloba Harvey (x2). (e) T. 

simmleri Beauverd (x4). (f) T. galpinii Schlechter (x2).


Plate III. (a) T. natalensis Baker (x3). (b) T. leucantha Baker (x2). (c) T. verdoorniae Vosa & Burbidge (x2). (d) T. coddiiVosa & 

Group D (five species) - The karyotype consists 

of five sub median chromosomes and one 

subterminal chromosome, usually the shortest 

of the complement, which carries the nucleolar 

organizer on its short am. The "satellite" distal 

to the nucleolar attachment is variable in size 

and in some cases may have almost the size of 

the arm of nucleolar chromosome type A. However, 

in all these cases, the extra chromosome 

segment is heterochromatic. 

Group E (one species) - The karyotype is 

similar to that of the species of group A, but the 

nucleolar attachment is carried on the long arm 

of a submedian chromosome.

104 VOSA 

Plate IV. (a) T. rhodesica Fries (x2). (b) T. violacea Harvey (x2). (c) T. violacea Harvey var. maritima Vosa (x4). (d) T. cominsiiVosa (x2). 

(e) The craggy hill of 'Unkomaninear Ofcolaco (Northern Drakensberg): from the top to the SW the habitat of T. transvaalensis Vosa. (f) 

Nabibia, small rocky inselberg about 75 km N of the Orange River with a population of Aloe dichotoma L. f: the typical habitat of T. 

tenuior Krause & Dinter.


The problem of the number of 

nucleolar organizers in Tulbaghia 

The number of nucleolar organizers in Tulbaghia 

vary from two to four and are of five 

types (fig. 22a). They differ in number and in 

combinations according to and characterizing 

the species (fig. 22b). 

Observation of silver-stained interphase nuclei 

in several species has shown that the amount of 

nucleolar material is very much equivalent 

whether there are two organizers or four. Thus, 

it seems that there has been an increase in the 

number of organizing sites with a proportional 

decrease in their nucleolar material and in its 

synthetizing activity. Silver-staining has also 

confirmed, in Tulbaghia, the intrinsic 

heteromorphism of the nucleolar organizers, 

and of their expression in the interphase nucleoli, 

which seems to be the rule in most organisms 

and based on a kind of dominance on the 

line of that demonstrated by NAVASHIN (1934) in 

hybrids between species of Crepis. 

The significance of the doubling of the nucleolar 

organizing sites must be found in the 

fine tuning of the interaction between the genome 

vis-a-vis the meiotic process and therefore 

on the inheritance and expression of certain 

adaptive features. The situation in T. violacea 

and T. cominsii, which share a very similar phenotype 

but with two organizers in the former 

and four in the latter, is rather interesting and 

demonstrates very well how little we known as 

yet on the very specialized biology and especially 

on the evolutionary meaning of such differences.

106 VOSA 

Constitutive heterochromatin in 

Tulbaghia 

Constitutive heterochromatin can be defined 

as somewhat dispensable, non-transcrib ing, 

usually highly repetitive DNA sequences, 

preserved by selection and located on the chromosomes 

as inheritable, inconvertible seg ments. 

It differs from facultative heterochromatin which 

is thought to be the consequence of a cytological 

situation in which a certain amount of normally 

coding DNA is temporarily and reversibly 

repressed by a special molecular mechanism. An 

example of facultative heterochromatin are the 

X-chromosomes in female mammals, one of 

which becomes inactive and heterochromatic 

and appears, as it were constitutive 

heterochromatin, as a chromocentre in the 

interphase nuclei (i.e. the Barr-body of female 

humans). 

Constitutive heterochromatic segments (Hsegments) 

are present in several species of Tulbaghia 

(Table 2 and see VOSA, 1970, 1975). The Hsegments 

number, size and position on the 

chromosomes is variable between and, to a certain 

extent, also within species. Such segments 

are sensitive to cold treatment and present reduced 

fluorescence when stained with certain 

fluorochromes, in the same way as those found 

in Paris, Trillium and Cestrum (DARLINGTON 

and LA COUR 1938; DYER 1963; VOSA 1966, 1970, 

1973; FILION and VOSA 1980). Experiments have 

shown that, in Tulbaghia, they consist mainly of 

highly repetitive sequences of DNA composed, 

for the most part, of Guanine-Cytidine nucleotides 

(PACHMAN and RIEGLER 1972; WEISBLUM and de 

HASETH 1972). 

In some species of Tulbaghia, such as T. cernua 

and T. ludwigiana, the total amount of heterochromatin, 

as a proportion of the length of 

the chromosome complement, has been found in 

a few plants to be as much as 25 %. 

In respect to the presence and to the localization 

of the H-segments, the species of Tulbaghia 

may be subdivided into three groups. 

The first group, which includes 10 species, the 

segments are generally very small and almost 

exclusively localized either side of the nucleolar 

organizers. The second group includes 6 spe cies 

where the H-segments are relatively large and 

are located almost always in distal position (fig. 

23). This situation is found also in most of the 

American species of Trillium (DARLINGTON and 

SHAW 1959). In the third group, which includes 5 

species, the H-segments are found mostly in 

intercalary position (fig. 24) as in Fritillaria, 

Trillium kamschaticum, Trillium erectum and 

Cestrum (DYER 1963). The variation in the 

presence, absence and size of the H-seg ments in 

Tulbaghia, follows a very similar pattern in 

different species within each group, with 

Fig. 23 — Mitotic metaphase in Tulbaghia cernua, after 36 h at 3-4 °C, stained in Feulgen (x 1500). Note the lightly stained distally 

located H-segments. The small segments, intercalary between the centromere and the N.O.'s, are also heterochromatic and lightly 

stained (arrowed).


Fig. 24 — C-banded mitotic metaphase in Tulbaghia leucantha (x 1500). Note the deeply stained intercalary H-segments. The chromosome 

portions, distal to the N.O.R.'s, are also heterochromatic and deeply stained (arrowed). 

an evident polymorphism which is almost exactly 

parallel for each chromosome type. This 

situation is very much the same as that found in 

Trillium and in Paris by DARLINGTON and SHAW 

(1959). In Trillium, in fact, the five basic chromosomes 

correspond in size and morphology 

throughout the American species, notwithstanding 

the position of the H-segments. In 

Tulbaghia, however, at least in the group with 

distal localization, the basic chromosome complement 

is variable according to the species. 

In our genus, the two main types of H-segments 

localization are distinct and only occasionally 

we have found intermediate types. In 

particular, in Tulbaghia leucantha, one of the 

species with proximally localized H-segments, a 

few plants in some populations have distally located 

segments of this type adjacent to the nu- 

TABLE 2 — Heterochromatin and B-chromosomes in Tulbaghia 

cleolar attachment. The heterochromatin position 

on the chromosomes does not affect the localization 

of chiasmata which are almost exclusively 

proximal to the centromere in all the species. 

As regards the H-segments found in 

Trillium, Fritillaria and in a few other genera, 

the general situation seems to suggest a clear 

evolutionary trend from an intercalary towards a 

distal localization (DARLINGTON and SHAW 1959). 

In Tulbaghia, the almost complete absence of 

intermediate types probably shows that the 

proximal localization of the H-segments may be 

of a more recent origin than the alternative, and 

that their establishment could be rather apart in 

time. The two situations are evidently adaptive in 

their function and in a state of finely balanced 

polymorphism within their respective 

interbreeding propulations. This problem has 

been estensively and masterly treated by FORD 

(1957). As regards Tulbaghia, a relatively small 

survey of heterochromatin polymorphism in two 

populations of T. leucantha (12 and 17 plants, 

respectively), has shown that the degree of 

chromosomal heterozygousity is quite high. 

Nevertheless, the same chromosome types have 

been found to be present in the same proportion 

in both samples, thus demonstrating that they 

belong to a single interbreeding population 

(VOSA 1973a). 

Function and significance of constitutive heterochromatin 

In Tulbaghia, as well as in many other genera 

of plants and animals, the presence of heterochromatin 

raises the question of its role in the 

framework of the genetic system. The function 

of heterochromatin seems to be overall nonspecific 

and quantitative rather than qualitative. 

A few studies have demonstrated the quantita-

108 VOSA 

tive function of heterochromatin. In Allium 

pulchellum, VOSA (1976, 1996) has shown that 

the differences in the amount of heterochromatin, 

between altitudinally separated populations, 

indicate a probable adaptive role. In Ornithogalum 

(VOSA 1997), the differences in the 

amount of heterochromatin are not only indicative 

of an adaptive role to extreme environmen tal 

conditions but, according to quantity, they have 

a strong influence in the sequence and du ration of 

the life cycle. 

The role of heterochromatin in the control of 

the life cycle has been studied by NAGL 

(1974a,b) and by NAGL and EHRENDORFER (1973). 

In the Anthemidae, NAGL (1974a,b) in his study 

of three perennial species, three annual species 

with high DNA content and two annual species 

with low DNA content, has found that the 

amount of heterochromatin per genome was 

directly related to a reduction in cell cycle time 

in the annual species with high DNA content. 

Now, because heterochromatin generally 

replicates much faster than euchromatin 

(BOSTOCK et al. 1972), its amount per genome 

may represent a regulating agent affecting the 

duration of the cell cycle and, therefore, of the 

speed of development. 

VOSA and STERGIANOU (1990), report some 

interesting results in their study of the genus 

Pleione (Orchidaceae), where there exist terrestrial 

as well as epiphytic species, all with the 

same chromosome number and equivalent DNA 

amount. Cytological data show that the 

terrestrial species generally possess asymmetric 

karyotypes and relatively small amounts of intestitial 

and centromeric heterochromatin 

whereas the epiphytic species have symmetric 

karyotypes and comparatively large amounts of 

mainly distally located heterochromatin. Thus, 

as in Anthemidae (NAGL 1974a,b) and in Ornithogalum 

(VosA 1997), the larger amount of 

heterochromatin may provide a kind of compensating 

factor which allows a quicker growth 

in the epiphytic species, which are known to 

complete their annual growing period in a rather 

shorter time than the terrestrial. Further, the 

differences in karyotype morphology and in the 

relative position of the H-segments in the two 

types of Pleione, may have an effect on meiosis 

and on crossing-over, with consequences in their 

respective heredity of gene sequences. In fact, a 

correlation between crossing-over and the 

presence of heterochromatin, 

and of its position on the chromosomes, has 

been reported in plants (RHOADES 1978) and in 

animals (MIKLOS and NANKIWELL 1976). 

In conclusion, constitutive heterochromatin 

owes its great evolutionary importance to its 

contribution to an overall increase in variation, 

which is probably not related to any decisive 

threshold but that controls both heredity and 

development. 

B-chromosomes in Tulbag/iia 

By definition B-chromosomes are smaller in 

size than those of the A-complement. They are 

somewhat unstable at mitosis and at meiosis and 

therefore variable in number between plants of 

the same populations. In general, Bchromosomes 

do not have any specific phenotypic 

consequence on the individual organism 

carrying them and seem to have quantitative 

rather than qualitative effects. Thus, they have 

been shown to increase the speed of germination 

in Allium porrum (VOSA 1966b), may have a 

role in adaptation to ecological situation in 

Listera ovata (VosA 1983) or may affect meiotic 

recombination in this latter species (VosA and 

BARLOW 1972) and in Puschkinia libanotica 

(BARLOW and VOSA 1970). The induced change in 

recombination, by creating new gene assemblages, 

may lead to speciation in the long term 

(DARLINGTON 1971). On the whole, the considerable 

amount of research has shown that Bchromosomes 

may be considered, with good 

reason, as the active basis of quantitative variation. 

In Tulbaghia, so far, only three species have 

been found to possess telocentric B-chromosomes 

(Table 2). In T. cernua, T. ludwigiana 

and T. cameronii (fig. 18), they are relatively 

large (3-4 micra in length) whilst in T. leucantha 

they are very small (1-2 micra in length). No 

population analysis regarding occurrence and 

breeding behaviour of B-chromosomes have 

been undertaken as yet in Tulbaghta. It is interesting 

that B-chromosomes are absent in the 

polyploid forms of T. cernua and T. leucantha, 

indeed a similar situation as that found in most 

plant species where, in general, their frequency 

has been found to be much less in polyploids 

than in diploids.


CONCLUSIONS 

The cytotaxonomic review of Tulbaghia has 

shown that the genus is fairly uniform with relatively 

small differences between the species. 

These differences relate chiefly to the size and 

type of the corona and to certain other morphological 

characters. 

However, the genus, as a whole, possesses 

large and well differentiated chromosomes which 

allow precise studies to be made on their 

structure. In particular, apart from specific differences 

in the size of the karyotypes and of the 

existence of heterochromatic segments, the type 

and number of the nucleolar chromosomes are 

data of great importance in the definition of the 

species and to subdivide the genus into groups 

of taxa which, with some exception, are 

composed of closely related species. 

In any case, for the recognition of the geographical 

limits of many species of Tulbaghia, 

more extensive field collecting trips are neces sary. 

Regarding South Africa, this applies particularly 

to the territories north of the rivers Vaal 

and Orange. 

A survey of the geographical distribution of 

the species of Tulbaghia, has shown than an apparent 

centre of speciation may be located in the 

Eastern Cape, where most of the species are 

found. The basis of this hypothesis may be not 

really valid because of the vegetation changes 

brought about by the climatic deterioration made 

worse, with every probability, also by human 

activities. 

These two factors seem to have been in action 

for at least the last 500-600 years. In particular, 

in quite recent times, the above mentioned 

climatic changes, together with instances 

of poor farm management and overgrazing, have 

been responsible for the advance southeastwards 

of desert-like conditions over a rather 

large area. This occurrence, which in some 

places amounts to about 250 km in width, according 

to ACOCKS (1953), has reduced drastically 

the habitat available for many plant species, 

especially in the north-west, leading in many 

instances to their disappearance. In the case of 

Tulbaghia, the net result has been an apparent 

"crowding" of the extant species in the east with 

a few somewhat specialized "relics", such as 

Tulbaghia tenuior and, in part, Tulbaghia 

dregeana, still able to survive in the west. 

Very few collections of Tulbaghia have been 

made north of the river Limpopo and, with a few 

exceptions, little is known about the tropi cal 

species of the genus. 

APPENDIX I 

Species insufficiently known 

Tulbaghia aequinoctialis Welwitsch, Baker 

in Trans. Linn. Soc. ser. II, 1: 146 (1878). Apparently 

found near Pungo Adongo in Angola. 

The type is at K (A.M. Reis 43-3754). I have 

seen two other specimens (Dinter 2140 and 

2280, BM!) but, in the absence of new evi dence, 

I consider the existing material not sufficient for 

an accurate appraisal of the epithet. 

Tulbaghia affinis Link in Enum. PI. Berol. p. 

310. Described as a plant from the Cape of 

Good Hope but without a precise locality. 

Tulbaghia australis Link in Steud. Nom. Bot. 

1: 857 (1821). It must be considered nomen nudum 

as no description is given and the name has 

been deleted from the second edition. 

Tulbaghia bragae Engler in Pflanzenw. Ost- 

Afrika, 141 (1895). Described from Braga near 

Beira (Mozambique). 

Tulbaghia calcarea Engler and Krause in 

Engl. Bot. Jarb. 45: 142 (1910). Described from 

around Grootfontein in Nabibia. On the basis of 

the criteria used in the present work, the existence 

of only two specimen in M (Dinter 761 

and 76la) is considered not sufficient to war rant 

its inclusion in the list of good species. 

Tulbaghia hockii de Wildemann in Feddes 

Rep. 11: 546 (1913). This entity has been described 

as from Upper Katanga (Elizabethville, 

Congo) but no specimens are indicated. 

Tulbaghia hypoxidea Smith in Rees Cycl. 36 

(1819). The description is apparently based on a 

plant that flowered in the greenhouses of 

Messrs. Lees & Kennedy of Hammersmith, 

London, who received it from Holland. A 

drawing of the plant was made by a Mr. Soverby 

but it is said to have been mislaid.

110 VOSA 

Tulbaghia luebbertiana Engler and Krause in 

Engl. Bot. Jahrb. 45: 145 (1910). Described from 

Nabibia but the type specimen seen, Luebbert 44 

(M!), is out of flower. 

Tulbaghia karasbergensis Glover in Ann. 

Bolus Herb. 1: 104 (1915). Although he has seen 

no material, Merxmueller, in Prod. Fl. 

Sudwestafrica of 1969, considers it a good species. 

APPENDIX II 

Taxonomy key to the genus Tulbaghia 

Tulbaghia monantha Engl. & Gilg. in Warb. 

Kunene-Zambesi Exp. 192 (1903). The species 

has been described as from Cabindere in Angola. 

The only specimen seen, Baum 351 (M!), in my 

opinion, cannot be taken as the basis of a distinct 

species. 

Tulbaghia pauciflora Baker in Bot. Jahrb. 

Syst. (1892). Described as from the Cape but no 

specimen is indicated.


Acknowledgements — I wish to thank the very 

many friends from various parts of the world, but especially 

from Africa, for their kindness and help during 

the collection and the cultivation of the material used 

in this study. 

I wish also to remember with deep gratitude, as 

the most helpful and knowledgeable colleagues and 

friends, the late Frank White M.A. for his general 

help in the taxonomic treatment, the late Prof. Cyril 

D. Darlington F.R.S. for his interest in my work, the 

late Prof. R.B. Rycroft Ph.D. and all the past and 

present Staff of the Kirstenbosch Botanic Garden for 

making available lodgings, transport and other essential 

facilities. I am very grateful too to Dr. David 

Mogford, D.Phil, and to Mr. Kenneth Burras, M.A., 

former Superintendent of the Botanic Garden of 

Oxford, for their help during some of the most important 

plant collecting trips. Very special thanks are 

also due to Prof. Emilio Battaglia for helpful discussions 

and appropriate comments on the manuscrit. 

REFERENCES 

ACOCKS J.P.H., 1953 — Veld types of South Africa. 

Mem. Bot. Surv. S. Afr. 28: 1-192. 

AVE-LALLEMANT J.E.L., 1844 — Tulbaghiae species quae 

hucusque innotuerunt expositae. Bull. Class Phys. 

Math. Acad. St. Petersb., 3: 1-11. 

BAKER J.G., 1871 —A revision of the genera and species 

of herbaceous capsular gamophyllous Liliaceae.Jout. 

Linn. Soc. 11: 349-436. 

BARLOW P.W. and VOSA C.G., 1970 — The effect of Bchromosomes 

on meiosis in Puschkinia libanotica. 

Chromosoma (Berl.), 30: 344-355. 

BOSTOCK C.J., PRESCOTT D.M. and HATCH F.T., 1972 — 

Timing replication of the satellite and main 

bandDNA in cells of the kangaroo-rat (Dipodomys 

ordii). Exp. Cell Res. 74: 487-495. 

BURBIDGE R.B., 1978 —A revision of the genus Tulhaghia 

(Liliaceae). Notes R. bot. Gdn. Edinb. 36: 

77-103. 

DARLINGTON C.D., 1971 — The evolution of polymorphic 

systems. In: "Ecological Genetics and Evolution", p. 

1-19. Blackwell, Oxford, London. 

DARLINGTON C.D. and LA COUR L.F., 1938 — Differential 

reactivity of the chromosomes. Ann. Bot. 2: 615- 

625. 

DARLINGTON C.D. and SHAW G.W., 1959 — Parallel 

polymorphism in the heterochromatin of Trillium 

species. Heredity, 13: 89-121. 

DYER A.F., 1963 — Allocyclic segments of chromosomes 

and the structural heterozygosity they reveal. 


FILION W.G. and VOSA C.G., 1980 — Quinacrine 

fluorescence studies in Paris polyphylla. Can. J. 

Genet. Cyt., 22: 417-420. 

FORD E.B., 1957 — Polymorphism in plants, animals 

and man. Nature, 180: 1315-1319. 

HOWELL W.M. and BLACK D.A., 1980 — Controlled 

silver-staining of Nucleolar Organizing Regions 

with a protective colloidal developer: a one-step 

method. Experientia, 36: 1014-1015. 

JACOBSEN J.U., YAMAGUCHI U., MANN L.K. and HOWARD 

F.D., 1968 — An alkyl-cysteine sulfox-ide lyase in 

Tulbaghia violacea and its relation to other 

alliinase-like enzymes. Phytochemistry, 7: 1099- 

1108. 

MIKLOS G.L. and NANKIWELL N., 1976 — Telomeric DNA 

function as regulating recombination. Chromosoma 

(Berl.), 56: 143-167. 

NAGL W., 1974a — Mitotic cycle in perennial and anual 

plants with varying amount of DNA and heterochromatin. 

Develpm. Biol. 39: 242-346. 

—, 1974b — Role of heterochromatin in the control of 

cell cycle duration. Nature, 248: 53-54. 

NAGL W. and EHRENDORFER F., 1973 — DNA-content, 

heterochromatin, mitotic index, and growth in 

perennial and annual Anthemidae (Asteraceae). PI. 

Syst. Evol. 123:35-44. 

NAVASHIN M., 1934 — Chromosome alterations 

caused by hybridization and their hearing upon 

certain general genetic problems. Cytology, 81: 

344-345. 

PACHMANN U. and RIEGLER R., 1972 — Quantum yield 

of acridine interaction with DNA of definite base 

sequences. A basis for explanation of acridine bands 

of chromosomes. Exp. Cell Res., 72: 602-608. 

RHOADES M.M., 1978 — Genetic effect of heterochromatin 

in Maize. In: WALDER D.B. (ed.) "Maize Breeding 

and Genetics", p. 641, Interscience, New York, 

U.S.A. 

UPHOF J.T., 1943 —A review of Agapanthus and Tulbaghia. 

Herbertia, 10: 40-51. 

VOSA C.G., 1966a — Chromosome variation in Tulbaghia. 

Heredity, 21: 395-412. 

—, 1966b — Seed germination and B-chromosomes in 

the Leek (Allium porrum). Chromosomes Today, 

1:24-27. 

—, 1966c — Tulbaghia hybrids. Heredity, 21: 675- 

687. 

—, 1970 — Heterochromatin recognition with fluorochromes. 

Chromosoma (Berl.), 30: 366-372, 

—, 1972 — Two-track heredity: differentiation of 

male and female meiosis in Tulbaghia. Caryologia, 

25:275-281. 

—, 1973 a — Quinacrine fluorescence analysis of chromosome 

variation in the plant Tulbaghia leucantha. 

Chromosome Today, 4: 345-349. 

—, 1973b — Heterochromatin recognition and analysis 

of chromosome variation in Scilla sibirica. 


—, 1975 — The cytotaxonomy of the genus Tulbaghia. 

Ann. Bot. (Rome), 34: 47-121.

112 VOSA 

—, 1976 — Heterochromatic banding patterns in Allium. 

II. Heterochromatin variation in species of 

the paniculatum group. Chromosoma (Berl.), 57: 

119-133. 

—, 1979 — Notes on Tulbaghia: 1. A new species 

from the Eastern Cape and a list of new localities. 

J.S. Afr. Bot.,45: 127-132. 

—, 1980 — Notes on Tulbaghia: 2. A. On the status of 

Tulbaghia violacea Haw. B. Tulbaghia fragrans 

Verdoom—Tulbaghia simmleri Beauv. C. Tulbaghia 

verdorniae Vosa &• Burbidge, two new localities. }. 

S. Afr. Bot. 46: 109-114. 

—, 1981 — Notes on Tulbaghia: 3. On the status of 

Tulbaghia cernua Ave-Lall. J. S. Afr. Bot. 47:57- 

61. 

—, 1982 — Notes on Tulbaghia: 4. A spontaneous hybrid 

from the Eastern Cape. J. S. Afr. Bot., 48: 

241-244. 

—, 1983 — Notes on Tulbaghia: 5. Scanning Electron 

Microscopy of the seed-coat patterns of nineteen 

species J. S. Afr. Bot., 49: 251-259. 

—, 1996 — Some aspects of karyotype evolution in 

Eiliflorae: heterochromatin variation and ecology 

in Alliumpulchellum. Bocconia, 5: 267-270. 

—, 1997 — Heterochromatin and ecological adaptation 

in Southern African Ornithogalum (Liliaceae). 

Caryologia, 50: 97-103. 

VOSA C.G. and BARLOW P.W., 1972 — The effects of Bchromosomes 

on meiosis in Listera ovata. Caryologia, 

25: 1-8. 

VOSA C.G. and STERGIANOU K., 1990 — The cytoecology 

of the genus Pleione (Orchidaceae). J. 

Orchid Society of India, 4: 29-35. 

WEISBLUM B. and DE HASETH P.L., 1972 — Quinacrine, a 

chromosome stain for deoxyadenylate-deoxythymidylate-rich 

regions in DNA. Proc. Nat. 

Acad. Sci. U.S.A., 69: 629-632. 

Received 4 April 2000; accepted 8 June 2000

A revised cytotaxonomy of the genus Tulbaghia (Alliaceae)

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