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FLORA OF SOUTHERN AFRICA

A taxonomic treatment of the flora of the Republic of South Africa, Lesotho, Swaziland and South-West Africa.

To be completed in 33 volumes, not in numerical sequence.

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BOTHALIA

Volume 12, No. 1

Edited by/Onder redaksie van

D. J. B. Killick

Editorial Committee/Redaksiekomitee: B. de Winter, D. Edwards, D. J. B. Killick and/en J. Ross

DEPT. YAK 1ANDB0P-TEBNIESE D1ENSTE

BIBLIOTEEK

NAV. IN8TITUUT V1R PLANTKUNDE

PRETORIA

BOTANICAL RESEARCH NSTITUTE

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1976

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CONTENTS-INHOUD

Volume 12, No. 1

Page

Bladsy

1. The South African species of Hemizygia (Lamiaceae). L. E. Codd 1

2. The genus Syncolostemon (Lamiaceae). L. E. Codd 21

3. Studies in the Ericoideae. I. The genera Eremia and Eremiella. E. G. H. Oliver 29

4. Studies in the Ericoideae. II. The new genus Stokoeanthus. E. G. H. Oliver 49

5. Notes on African plants

Asclepiadaceae. R. A. Dyer 53

Blechnaceae. J. C. Scheepers and P. Vorster 57

Ericaceae. E. G. H. Oliver 57

Fabaceae. J. H. Ross 59

Liliaceae. J. P. Jessop; A. A. Obermeyer; D. S. Hardy 60

Melastomataceae. J. H. Ross 62

Mesembryanthemaceae. H. R. Tolken and J. P. Jessop 63

6. A procedure for standardizing comparative leaf anatomy in the Poaceae. I. The leaf blade as viewed

in transverse section. R. P. Ellis 65

7. Studies in the Hypoxidaceae. I. Vegetative morphology and anatomy. M. F. Thompson Ill

8. Notes on Veronaea including V. compacta sp. nov. M. C. Papendorf 119

9. The soil mycoflora of an Acacia karroo Community in the Western Transvaal. M. C. Papendorf 123

10. The mycoflora of wheat field debris, Part II. W. J. Jooste 129

11. Hans Joachim Schlieben, collector extraordinary. O. A. Leistner 133

12. A phytosociological classification of the Nylsvley Nature Reserve. B. J. Coetzee, F. van der

Meulen, S. Zwanziger, P. Gonsalves and P. Weisser 137

13. Thuranthos: notes on generic status, morphology, phenology and pollination biology. C. H. Stirton 161

Book Reviews 167

-

Bothalia 12, 1: 1-20 (1976)

The South African species of Hemizygia (Lamiaceae)

L. E. CODD*

ABSTRACT

The South African species of Hemizygia are reviewed and 28 species are recognized, including the following

new names: H. macrophylla (Guerke) Codd ( =SyncoIostemon macrophyllus Guerke), H. pretoriae Guerke var.

heterotricha Codd, H. cinerea Codd, H. incana Codd, H. modesta Codd, H. parvifolia Codd, H. punctata Codd

and H. ramosa Codd.

Resume

LES ESPECES SUD-AFRICAINES D’HEMIZYGIA ( LAMIACEAE )

Les especes sud-africaines c/'Hemizygia sent examinees et 28 especes sont reconnues y compris les noms

nouveaux suivants: H. macrophylla ( Guerke ) Codd ( Syncolostemon macrophyllus Guerke), H. pretoriae Guerke

var. heterotricha Codd, H. cinerea Codd, H. incana Codd, H. modesta Codd, H. parvifolia Codd, H. punctata

Codd et H. ramosa Codd.

INTRODUCTION

Hemizygia was initially proposed by Bentham as

a section of Ocimum in DC., Prodr. 12:41 (1848), to

accommodate the single species O. teucriifolium

Hochst., in which the filaments of the lower (anticous)

pair of stamens are fused at the base, in contrast to

the free filaments of typical Ocimum species. Briquet

raised the section to generic rank in Pflanzenfam.

4,3a:368 (1897) and subsequently described several

species. He laid stress on the fused filaments of the

anticous stamens, in which respect it resembled

Syncolostemon E. Mey. ex Benth., but differed from

the latter in the broadly ovate upper tooth of the

calyx.

Baker in FI. Trop. Afr. 5:365 (1900) and N. E.

Brown in FI. Cap. 5, 1 :237 (1910) included Hemizygia

in Orthosiphon, the latter author pointing out that

the union of the filaments may vary in some species

and even on the same plant.

Ashby in J. Bot. Lond. 73:312, 343 (1935) resus-

citated the genus Hemizygia, remarking that although

the degree of union of the filaments varies, only

very rarely are they free to the base. He also noted

other differences in the androecium of Hemizygia

and Orthosiphon. The anticous pair of stamens in

Hemizygia are contiguous at their insertion at the

mouth of the corolla tube, while in Orthosiphon

they are well separate at their insertion, which is at a

short distance within the corolla tube, or rarely in

the throat. The posterior stamens are inserted from

about the middle to near the base of the corolla tube

in Hemizygia (with one exception, H. gerrardii) and

are well exserted (with the exceptions of H. pretoriae

and H. persimilis), while in typical Orthosiphon the

upper stamens are inserted near the throat and are

only shortly exserted. It may also be noted that in

Hemizygia the filaments are usually pubescent in the

lower part, but in typical Orthosiphon they are

glabrous.

He also drew attention to certain differences in

corolla structure between the two. In Hemizygia the

corolla tube is usually dilated at the throat and

truncate at the mouth; the posterior lip is small and

* Botanical Research Institute, Department of Agricultural

Technical Services, Private Bag X101, Pretoria.

the anterior lip larger and often deflexed at maturity

In contrast, typical Orthosiphon rarely has the corolla

tube widened at the throat, the mouth is not truncate

but the two lips meet laterally and are about equal in

length. He concludes that the exceptions are suffi-

ciently few that they should not necessitate the merging

of Hemizygia in Orthosiphon.

This view is supported in the present study and

reference may be made to my review of the South

African Orthosiphon species in Bothalia 8:149 (1964).

Here it was noted that the species separated by

Bremekamp in his genus Nautochilus, together with

those of Orthosiphon sect. Serrati Ashby, form an

aberrant group, which was placed as Orthosiphon

subgen. Nautochilus (Brem.) Codd. In these species

the filaments of the posterior stamens are attached

near the base of the corolla tube and are pubescent,

as in Hemizygia, while the anterior filaments are

free to the base, as in Orthosiphon. The corolla shape

in these species, although somewhat intermediate,

agrees better with Orthosiphon than with Hemizygia.

Differences between Hemizygia and Syncolostemon

must also be found if the former genus is to be upheld,

and these are discussed in another article (Bothalia

11: 21 1976). The corolla and androecium characters

are virtually the same in the two genera and it is

mainly the calyx which provides a basis for distin-

guishing them. In Hemizygia the upper calyx tooth is

broadly ovate and decurrent on the tube while the

lower 4 teeth are usually subulate to spinescent; in

typical Syncolostemon, on the other hand, the calyx

is subequally 5-toothed. Two intermediate species

were noted in which the upper tooth was elliptic or

broadly elliptic, not decurrent on the tube. These

were included in Syncolostemon because of their

close affinity with S. rotundifolius and S. densiflorus.

Orthosiphon, Syncolostemon and Hemizygia form a

closely related series of which Orthosiphon (1830) is

the oldest name. The great majority of species can

be allocated without great difficulty to one of the

three genera now upheld and this appears to be

sufficient justification for continuing with their

present circumscription.

A striking feature of some Hemizygia species is the

strong development of the apical bracts of the

inflorescence into a persistent colourful tuft or coma.

2

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

The genus Bouetia A. Chev. in Mem. Soc. Bot. Fr.

2:200 (1917) was based on such a species, B. ocimoides

A. Chev., which is generally regarded as a synonym

of Hemizygia bracteosa (Benth.) Briq.

HEMIZYGIA

Hemizygia {Benth.) Briq. in Pflanzenfam. 4,3a:368

(1897); Annu. Conserv. Jard. Bot. Geneve 2:247

(1898); Ashby in J. Bot. Lond. 73:312,343 (1935);

Phillips, Gen. ed. 2:652 (1951); Compton, FI. Swaz.

67,158 (1966); Launert & Schreiber in Prodr. FI.

S.W. Afr. 123: 11 (1969); Ross, FI. Natal 306 (1972).

Type species: H. teucriifolia (Hochst.) Briq.

Orthosiphon sensu Bak. in FI. Trop. Afr. 5: 365

(1900), partly; sensu N.E. Br. in FI. Cap. 5,1;237

(1910), partly.

Bouetia A. Chev. in Mem. Soc. Bot. Fr. 2:200

(1917). Type species: B. ocimoides A. Chev.

Perennial soft shrubs or annual herbs, or stems

arising annually from a perennial woody rootstock.

Leaves sessile or petiolate, variously pubescent and

gland-dotted, usually toothed. Inflorescence usually

of terminal racemes, lax or dense; verticillasters 2-6-

flowered; bracts small or the terminal ones enlarged

and persisting as a colourful coma. Calyx with the

upper tooth the largest, broadly ovate to subrotund,

decurrent on the tube; lower 4 teeth subulate to

spinescent, rarely deltoid-lanceolate. Corolla bila-

biate; tube longer than the calyx, widening from about

the middle to a truncate mouth and sometimes slightly

narrowed again at the mouth; upper lip small,

usually much shorter than the lower lip; lower lip

concave, horizontal to deflexed. Stamens 4, exserted

(upper pair included in H. pretoriae), didynamous;

upper pair affixed about or below the middle of the

corolla tube (above the middle in H. gerrardii),

filaments free, usually pubescent below and

occasionally higher; lower pair attached at the

corolla throat, filaments connate for part or the

entire length (occasionally almost free), glabrous.

Style exserted, usually minutely bifid, occasionally

clavate. Nutlets ovoid.

Key to Species

Stellate (branched) hairs present on leaves and other parts, often intermingled with simple hairs:

Upper (posterior) stamens included in the corolla tube 15b. H. pretoriae subsp. heterotricha

Upper (posterior) stamens exserted from the corolla tube, usually well exserted:

Leaf margin flat, not revolute:

Leaf margin crenate-dentate (sometimes indistinct in H. incana, H. cinerea and H. parvifolia ):

Verticillasters 3-6-flowered :

Leaf blade 5-9 cm long; inflorescence laxly branched, up to 60 cm long; calyx setose

in the throat 1. H. macrophylla

Leaf blade less than 5 cm long (rarely to 6 cm in H. obermeyerae ); inflorescence lax to

dense, up to 25 cm long; calyx not setose in the throat:

Leaf blade ovate, 2,5-6 cm long, subglabrous and rugose above. .2. H. obermeyerae

Leaf blade lanceolate to narrowly elliptic, grey-velvety above:

Calyx 8-9 mm long; corolla 12-15 (tube 10-12) mm long; leaves 15-35 x

6-12 mm, upper surface coarsely velvety 5. H. incana

Calyx 5-7 mm long; corolla 8-11 (tube 6-9) mm long; leaves 7-20 x 2-7 mm,

upper surface finely velvety, often darker than the lower 6. H. cinerea

Verticillasters 2-flowered :

Leaf blade small, rarely exceeding 2,5 cm long, upper surface subglabrous, rugose;

petiole up to 2 mm long:

Leaf blade lanceolate-elliptic, 15-25 mm long; apical bracts conspicuous, up to

15 mm long 3. H. rugosifolia

Leaf blade ovate, 6-11 mm long; apical bracts inconspicuous, up to 3,5 mm

long 4. H. parvifolia

Leaf blade ovate, usually exceeding 2,5 cm long, sparsely to densely floccose on

both surfaces; petiole 5-8 mm long, densely floccose 9. H. floccosa

Leaf margin entire:

Leaf blade lanceolate-elliptic to linear-elliptic, upper side often darker and with finer

tomentum; bracts inconspicuous, about 5 mm long 6. H. cinerea

Leaf blade ovate-lanceolate to broadly ovate, densely grey velvety on both surfaces;

bracts colourful, 7-10 mm long:

Calyx 5 mm long; upper stamens pubescent to above the middle; stigma

capitate 7. H. elliottii

Calyx 8-10 mm long; upper stamens pubescent only in the lower part; stigma shortly

bifid 8. H. gerrardii

Leaf margin revolute:

Corolla tube widening towards the mouth; stamens exserted well beyond the anterior

lip of the corolla:

Bracts 10-15 mm long, colourful 10. H. stenophylla

Bracts up to 7 mm long, inconspicuous:

Leaves finely grey velvety on both surfaces; stem finely grey tomentulose. .6. H. cinerea

Leaves coarsely stellate-pubescent and often yellowish below, much darker and

rugose above; stems villous 11. H. rehmannii

Corolla tube cylindrical, often slightly narrowed at the mouth; stamens exserted scarcely

beyond the anterior lip of the corolla:

Lower internodes of main stems usually more than 2 cm long; leaves 3-6 mm or

more broad, especially the lower 13. H. teucriifolia

Lower internodes of main stems less than 2 cm long; leaves usually not more than

4 mm broad 12. H. subvelutina

L. E. CODD

3

Stellate (branched) hairs absent:

Leaves narrow, leathery, revolute at the margin, thickly tomentose beneath with long white

hairs, somewhat varnished above 14 // albiflora

Leaves broad or narrow, not revolute at the margin, glabrous to tomentose beneath but not

as above:

Terminal bracts of the inflorescence like the lower ones, deciduous, small and incon-

spicuous :

Upper (posterior) stamens included in the corolla tube. ..15a. H. pretoriae subsp. pretoriae

Upper (posterior) stamens exserted from the corolla tube:

Verticillasters 2-flowered; leaves 6-15 x 3-7 mm:

Stems 12-25 cm tall, usually sparingly branched, arising annually from a

woody rootstock 16. H. modesta

Stems 60-120 cm tall, shrubby, much branched 17. H. punctata

Verticillasters 4— 6-flowered, leaves usually longer than above:

Leaves elliptic-ovate to broadly ovate, obtuse to rounded at the apex, obtuse

or broadly cuneate at the base; stems 25-40 cm arising annually from a

woody rootstock 18. H. bolusii

Leaves linear to ovate, apex acute, base cuneate; annual or perennial herbs not

arising from a perennial woody rootstock:

Stem and leaves with pubescence of short or fairly short dense and often

crisped hairs :

Leaves ovate-lanceolate to ovate; petiole 6-14 mm long. . . .24. H. petiolata

Leaves linear to lanceolate or, rarely, ovate-lanceolate; petiole usually

less than 5 mm long 25. H. canescens

Stem villous to subglabrous, not as above; leaves subglabrous or sparingly

pubescent to canescent or villous, often with long and short hairs inter-

mingled :

Leaves linear or with some leaves on a plant up to 5 mm broad, sub-

glabrous; stems subglabrous with few long hairs, often somewhat

varnished 26. H. linearis

Leaves linear-lanceolate to ovate-lanceolate, usually more than 5 mm broad;

stems and leaves sparingly to densely villous 27. H. petrensis

Terminal bracts of inflorescence distinct from the lower ones, membranous, forming a

persistent colourful coma (often small but coloured in H. petiolata and H. petrensis) :

Stamens not exserted beyond the lower lip of the corolla; filaments of upper pair of

stamens pubescent from the base to near the apex 23. H. persimilis

Stamens exserted beyond the lower lip of the corolla; filaments of upper pair

pubescent only near the base:

Verticillasters 2-flowered:

Stems shrubby, up to 1 m tall, much-branched ; leaves obovate to oblanceolate,

15-25 X 6-11 mm; corolla 25-28 mm long 19. H. ramosa

Stems up to 30 cm long arising annually from a woody rootstock; leaves ovate,

usually exceeding 25 mm long and 11 mm wide; corolla 12-15 mm long

21. H. foliosa

Verticillasters 4-6-flowered :

Terminal bracts ovate to linear-lanceolate, cuneate at the base, pairs of bracts

often spaced 1-2 cm apart, more than twice as long as broad (sometimes

less in H. transvaalensis but then corolla tube more than 12 mm long):

Corolla tube more than 12 mm long; terminal bracts ovate to lanceolate,

rarely linear-lanceolate 20. H. transvaalensis

Corolla tube less than 12 mm long; terminal bracts lanceolate to linear-

lanceolate 22. H. thorncroftii

Terminal bracts broadly ovate, not cuneate at the base, densely crowded,

usually less than twice as long as broad:

Petiole of mature leaves more than 5 mm long; leaves covered beneath with

a fine greyish-white pubescence 24. H. petiolata

Petiole of mature leaves less than 5 mm long or leaves sessile; underside of

leaves glabrous to variously pubescent:

Terminal bracts conspicuous, violet or white, up to 14 x 9 mm; corolla

white 28. H. bracteosa

Terminal bracts small, often purplish, about 5x3 mm; corolla mauve

27. H. petrensis

1. Hemizygia macrophylla ( Guerke ) Codd, comb.

nov.

Syncolostemon macrophyllus Guerke in Bull. Herb. Boiss.

6: 555 (1898); Ross, FI. Natal 306 (1972). Type: Natal, Drakens-

berg, Rehmann 7016 (Z, holo.).

Orthosiphon macrophyllus (Guerke) N. E. Br. in FI. Cap.

5, 1:241 (1910).

Soft shrub 1-1,5 m tall, branching from the base,

highly aromatic; stems several, woody at the base,

arising from a perennial woody rootstock, sparingly

branched, leafy towards the base, densely grey

pubescent. Leaves shortly petiolate; petiole up to

4 mm long; blade ovate-lanceolate to lanceolate,

6-9 cm long, 2,5-3 cm broad, both surfaces densely

and coarsely stellate-velvety, tending to fold along

the midrib and then somewhat falcate; apex acute,

base cuneate, tapering into the petiole; nerves impres-

sed above, reticulate beneath; margin regularly serrate

in the upper two-thirds. Inflorescence very lax, branch-

ed, up to 60 cm long and 25 cm broad; rhachis gland-

ular-hispid; bracts early caducous, broadly ovate,

acute, 6-8 mm long, concave, with a white margin and

patches of white tomentum; verticillasters 3-6-

flowered, up to 2 cm or more apart. Calyx 7 mm long,

glandular-hispid, setose in the throat, becoming

swollen and narrow at the mouth when in the fruiting

stage; upper lip ovate, acuminate, 2 mm long: lower

4 teeth deltoid-subulate, 2 mm long. Corolla purple,

17-18 mm long, pubescent on the lobes; tube 11-

12 mm long, widening to 5-6 mm at the mouth;

upper lip a small appendage, 1 mm long; lower lip

concave, 5-6 mm long, horizontal. Stamens exserted

4

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

well beyond the lower lip, curled upwards; upper

pair attached below the middle of the corolla tube,

puberulous near the base; lower pair united to near

the apex. Stigma bifid.

Recorded from the foothills of the Drakensberg

in northern Natal and southern Transvaal, in dense

grass, often among dolerite rocks, at altitudes from

1 500 to 1 800 m. Flowering is mainly from January

to April.

Transvaal. — 2730 (Vryheid): Mooihoek (-BC), Devenish 444

Natal. — 2729 (Volksrus): Normandien Pass (-DC), Codd

9979; near Ingogo (-DB), Medley Wood 6398 (K); Obermeyer

sub TR V 35941 ; Codd 9694; Mayne s.n.

Distinguished from all other species by the large

leaves covered on both surfaces with a dense, coarse

stellate pubescence and by the large, laxly branched

inflorescence. Its nearest affinity is probably with

Syncolostemon parviflorus but the upper calyx tooth

is broadly ovate. It is, therefore, somewhat inter-

mediate between Hemizygia and Syncolostemon.

2. Hemizygia obermeyerae Ashby in J. Bot. Lond.

73: 343 (1935). Type: Soutpansberg, Entabeni,

Obermeyer sub TRV 31556 (PRE, holo.).

Soft shrub, freely branched, 1-1,5 mm tall;

branches shortly stellate tomentose. Leaves petiolate;

petiole 6-12 mm long; blade broadly ovate to ovate-

lanceolate, 3-6 cm long, 1,8-3 cm broad, sub-

glabrous, brownish and rugose above, fairly densely

grey stellate pubescent below; apex obtuse to rounded,

base truncate to obtuse; margin finely crenate-

dentate. Inflorescence usually branched, fairly dense,

8-18 cm long, up to 10 cm in diameter; rhachis

stellate pubescent; bracts persisting at the apex,

mauve-purple, ovate, acute, up to 10-15x5-10 mm,

sparingly pubescent; verticillasters 6-flowered, 1-2 cm

apart. Calyx 8 mm long, glandular-setulose; upper

lip ovate, 2 mm long, decurrent on the tube; lower

4 teeth deltoid-subulate, the lowest pair the

longest, 2 mm long. Corolla mauve-pink, 18-22 mm

long, glabrous; tube 15-17 mm long, widening to

6-8 mm at the mouth; upper lip a small appendage,

1 mm long: lower lip concave, 4-6 mm long,

horizontal, to slightly deflexed. Stamens exserted well

beyond the lower lip, curled upwards; upper pair

attached below the middle of the tube, puberulous at

the base; lower pair united to the apex. Stigma bifid.

Grows with bracken and shrub on stony hillsides

and forest margins at altitudes of 1 400 to 1 800 m

in north-eastern Transvaal.

Transvaal. — 2230 (Messina): Entabeni Forest Reserve

(-CC), Obermeyer sub TRV 31556 ; 876; Bruce <t- Kies 8 A;

Taylor 727 ; Codd 4188; 8393; Piesanghoek (-CC), Gerstner

5746\ Pepiti Falls (-CD), Smuts & Gillett 3189; near Lake

Funduzi (-CD), Weintroub sub J35682; Thate Vondo Forest

Reserve (-CD), Van Graan & Hardy 564. 2330 (Tzaneen):

Woodbush (-CC), Hutchinson 2238. 2430 (Pilgrim’s Rest):

The Downs (-AA), Codd 9472; Marais 96; near The Downs

(-AC), Vahrmeijer 2360.

A distinct species easily separated from others

with stellate pubescence by the large, petiolate leaves.

With its large purplish bracts and mauve-pink

flowers, this is a showy species which grows well under

humid conditions but has not succeeded in cultivation

in the drier parts of the Transvaal.

3. Hemizygia rugosifolia Ashby in J. Bot. Lond.

73: 344 (1935). Type: Transvaal, The Downs, Junod

4342 (PRE, holo.).

Erect soft shrub, branched, probably about 1 m

tall; branches shortly stellate tomentose. Leaves

shortly petiolate; petiole 2-4 mm long; blade ovate-

lanceolate to elliptic, 15-25 mm long, 6-10 mm broad,

slightly coriaceous, upper surface rugose, puberulous

and with nerves immersed, lower surface densely and

shortly greyish stellate pubescent; apex obtuse, base

cuneate; margin finely and regularly crenate-dentate.

Inflorescence usually branched, medium lax, 8-13 cm

long; rhachis puberulous and gland-dotted; bracts

persisting at the apex, purplish, ovate, acute to

acuminate, about 10x5 mm, subglabrous with a

fringe of hairs; verticillasters 2-flowered, 1-1,5 cm

apart. Calyx 10 mm long, glandular-hispidulous;

upper lip ovate, 3 mm long, decurrent on the tube;

lower 4 teeth deltoid-subulate, the lowest pair much

longer than the median, narrowly subulate, 3 mm

long. Corolla 22 mm long; tube 18 mm long, widening

to 5-6 mm at the mouth; upper lip a small appendage

1 mm long; lower lip concave, 4 mm long, usually

deflexed. Stamens well exserted beyond the lower lip;

lower pair united to near the apex. Stigma minutely

bifid.

Known only from three gatherings near The Downs

in north-eastern Transvaal, where it apparently

grows at forest margins.

Transvaal.— 2430 (Pilgrim’s Rest): The Downs (-AA),

Junod 4342; Rogers 20188; Crundall s.n.

A small-leaved species related to the two species

described below, H. parvifolia and H. cinerea. From

H. parvifolia, which also has 2-flowered verticillasters,

it differs in the longer, more lanceolate-elliptic leaves

and the larger apical bracts; H. cinerea also has

small bracts and 6-flowered verticillasters but the

leaves, which are similar in shape to H. rugosifolia,

are finely grey velvety pubescent on both sides.

H. rugosifolia is a little known species last collected

in 1945.

4. Hemizygia parvifolia Codd, sp. nov., a H.

rugosifolia Ashby foliis parvioribus, ovatis, inflores-

centiis brevioribus, bracteis parvioribus differt.

Frutex, ramosus, 50-100 cm altus; ramuli stellato-

floccosi. Folia breviter petiolata; petiolus 1-2,5 mm

longus, dense stellato-floccosus; lamina ovata vel late

ovata, 6-11 mm longa, 4-9 mm lata, discolor, supra

rugosa, brunnea, hispidula vel subglabra, subtus

cinerea, dense stellato-tomentosa, nervis supra im-

pressis, subtus reticulatis, apice obtuso vel rotundato,

basi obtusa vel truncata, margine minute crenato-

dentato. Inflorescentia simplex vel basin versus parce

ramosa, 5-8 cm longa; rhachis stellato-floccosa ;

bracteae ovatae, 2, 5-3, 5 mm longae, caducae, basin

versus stellato-pubescentes; verticillastri 2-flori; pedi-

celli 3-4 mm longi, stellato-hispidi. Calyx 9-10 mm

longus, stellato-hispidus, glanduloso-punctatus; tubus

6 mm longus; lobus posticus late ovatus, acutus,

suberectus, 2-3 mm longus, margine decurrente;

dentes laterales deltoideo-subulati, 1,5 mm longi;

antici subulati, 2,5 mm longi. Corolla alba, 15-17 mm

longa, glabra vel extus labiis puberulis; tubus 11-14

mm longus, rectus, apicem versus sensim ampliatus,

ore 4 mm lato; labium posticum parvum; anticum

concavum, 4-5 mm longum. Stamina 12-14 mm

exserta; postica circa medium tubi corollae inserta,

filamentis liberis, prope basin pubescentibus; antica

fauce corollae inserta, filamentis omnino ad apicem

connatis. Stylus 10-12 mm exsertus, apice breviter

bilobato.

Type. — Transvaal, 2430 (Pilgrim’s Rest), farm

Belvedere, overlooking Blyde River Gorge (-DB),

Codd 10321 (PRE, holo.).

Shrub, branched, 50-100 cm tall; branchlets stellate-

floccose. Leaves shortly petiolate; petiole 1-2,5 mm

long, densely stellate-floccose; blade ovate to broadly

ovate, 6-1 1 mm long, 9 mm broad, discolorous, upper

surface rugose, brown, hispidulous to subglabrous,

L. E. CODD

5

lower surface grey, densely stellate-tomentose, nerves

impressed above, reticulate below, apex obtuse to

rounded, base obtuse to truncate, margin minutely

crenate-dentate. Inflorescence simple or sparingly

branched towards the base, 5-8 cm long; rhachis

stellate-floccose; bracts ovate, 2, 5-3, 5 mm long,

caducous, stellate-pubescent towards the base; verti-

cillasters 2-flowered; pedicels 3-4 mm long, stellate-

hispid. Calyx 9-10 mm long, stellate-hispid and

gland-dotted; tube 6 mm long; posticous lobe

broadly ovate, acute, suberect, 2-3 mm long, margin

decurrent; lateral teeth deltoid-subulate, 1,5 mm

long; anticous teeth subulate 2,5 mm long. Corolla

white 15-17 mm long, glabrous with outer surfaces

of lips puberulous; tube 11-14 mm long, straight,

widening gradually towards the apex, mouth 4 mm

wide; posticous lip small; anticous lip concave,

4-5 mm long. Stamens exserted by 12-14 mm; posticous

stamens inserted about the middle of the corolla tube

filaments free, pubescent near the base; anticous

stamens inserted in the corolla throat, filaments

completely united to the apex. Style exserted by 10-12

mm, apex shortly bilobed. Fig. 1.

Fig. 1. — Hemizygia parvifolia ( Codd 9802, PRE, holotype).

Found among quartzite rocks at altitudes of 1 300

to 1 500 m on the eastern Transvaal Drakensberg

escarpment; in flower from October to March.

Transvaal.— 2430 (Pilgrim’s Rest): 6 km N. of Vaalhoek

(-DB), Codd 9802; farm Belvedere, overlooking Blyde River

Gorge (-DB), Codd 10321; Blyde River Reserve (-DB), Davidson

2663; Blyde River hills (-DB), Ranh & Schlieben 9689 ; Bourke’s

Luck (-DB), Davidson & Mogg 33354; Davidson 73 (J); 2530

(Lydenburg): Nelshoogte Forestry Station, “The Knuckles ’

(-DD), Codd 9555.

H. parvifolia is a much-branched shrub up to 1 m

tall, related to H. rugosifolia Ashby, a species known

as yet only from The Downs, grid 2430 (-AA), some

100 km to the north-west of Blyde River Gorge, on the

same escarpment. H. parvifolia may be separated on

the basis of the smaller, more broadly ovate leaves,

the grey indumentum of the underside of the leaves,

the smaller inflorescence which is simple or sparingly

branched, the shorter bracts and the presence of

stellate hairs on the calyx.

It would be interesting to know whether either

species or intermediates occur on the escarpment

between The Downs and Blyde River Gorge. This is a

relatively inaccessible area which is scarcely known

botanically. Further study is also required of the

escarpment between the Blyde River Gorge and the

other known locality on the Nelshoogte Forestry

Station near Kaapsche Hoop.

5. Hemizygia incana Codd, sp. nov, a H. rugosifolia

Ashby foliis dense griseo-tomentosis, bracteis par-

vioribus, calyce villoso differt.

H. rehmannii sensu Ashby in J. Bot. Lond. 73: 347 (1935),

pro parte quoad Wager sub TRV 15598.

Frutex parce ramosus, 60 cm altus; ramuli dense

griseo-tomentosi, pilis stellatis et longis simplicibus.

Folia sessilia vel subsessilia; lamina ovata vel

lanceolata vel elliptico-lanceolata, 1,5-3, 5 cm longa,

6-12 mm lata, dense stellato-velutina, subtus argentea,

supra cinerea, nervis obscuris, apice obtuse vel

rotundato, basi obtusa, margine supra medium

minute crenato-dentato. lnflorescentia simplex vel

basin versus parce ramosa, 8-20 cm longa; rhachis

dense albo-tomentosa; bracteae late ovatae, acumina-

tae, caducae, parce vel dense pubescentes; verti-

cillastri plerumque 6-flori; pedicelli 2-3 mm longi

villosi. Calyx 8-9 mm longus, purpureo-suffusus,

glanduloso-villosus; tubus 5 mm longus; lobus

posticus late ovatus, acutus, suberectus, 2-3 mm

longus, margine decurrente; dentes laterales deltoideo-

subulati, 1,5 mm longi; antici subulati 2,5 longi.

Corolla malvina, 12-15 mm longa, glabra vel extus

labiis puberulis; tubus 10-12 mm longus, rectus,

apicem versus sensim ampliatus, ore 3-4 mm lato;

labium posticum 1,5 mm longum; anticum con-

cavum, 3 mm longum. Stamina 10-12 mm exserta;

postica prope basin tubi corollae inserta, filamentis

liberis prope basin pubescentibus; antica fauce

corollae inserta, filamentis fere ad apicem connatis.

Stylus 15 mm exsertus, apice breviter bifidus.

Type: Transvaal, 2530 (Lydenburg), Kaapsche

Hoop (-DB), Codd 5758 (PRE, holo.).

Shrub, sparingly branched, 60 cm tall; branchlets

densely grey-tomentose with stellate and long simple

hairs. Leaves sessile to subsessile; blade ovate or

lanceolate to elliptic-lanceolate, 1,5-3, 5 cm long,

6-12 mm broad densely stellate-velvety, silvery

below, darker grey above, nerves obscure, apex

obtuse to rounded, base obtuse, margin minutely

crenate-dentate above the middle. Inflorescence simple

or sparingly branched towards the base, 8-20 cm

long; rhachis densely white-tomentose; bracts broadly

ovate, acuminate, caducous, sparingly to densely

pubescent; verticillasters usually 6-flowered; pedicels

2-3 mm long, villous. Calyx 8-9 mm long, purple-

tinged, glandular-villous with long white hairs and

short gland-tipped hairs; tube 5 mm long; posticous

lobe broadly ovate, acute, suberect, 2-3 mm long,

margin decurrent; lateral teeth deltoid-subulate,

1,5 mm long; anticous teeth subulate, 2,5 mm long.

Corolla mauve, 12-15 mm long, glabrous, with the

outer surfaces of the lips puberulous; tube 10-12 mm

6

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

long, straight, widening gradually towards the apex,

mouth 3-4 mm wide; posticous lobe 1,5 mm long;

anticous lobe concave 3 mm long. Stamens exserted

by 10-12 mm; posticous stamens inserted near the

base of the corolla tube, filaments free, pubescent

near the base; anticous stamens inserted in the

throat of the corolla, filaments united almost to the

apex. Style exserted by 15 mm, apex shortly bifid.

Fig. 2. '

Fig. 2. — Hemizygia incana ( Codd 5758, PRE, holotype).

Found in shallow sandy soil among quartzite

rocks in the neighbourhood of Kaapsche Hoop at an

altitude of about 1 800 m; flowering has been recorded

from September to January.

Transvaal. — 2530 (Lydenburg): Berlin, Godwan River

(-DA), Hofmeyr 98; Kaapsche Hoop (-DB), Wager sub TRV

15598; Pole Evans 984; Thode A 1637; Codd 5758.

H. incana is related to H. rugosifolia Ashby but

may readily be distinguished by the dense grey-white

tomentum on both surfaces of the leaves, obscuring

the veins, the smaller and more pubescent bracts and

the villous calyx. For differences between this species

and H. cinerea Codd, described below, see notes

at the end of the latter description.

6. Hemizygia cinerea Codd, sp. nov., a H. incana

Codd foliis parvioribus, cinereis, floribus parvioribus

differt.

El. elliottii sensu Ashby in J. Bot. Lond. 73: 345 (1935),

pro parte quoad Bayer & McClean 217; Hutchinson, Forbes

& Verdoorn 52; sensu Ross, FI. Natal 306 (1972), pro parte

quoad spec. cit.

H. stenophylla sensu Ashby, l.c. 347 (1935), pro parte quoad

Galpin 10168; 11846; sensu Edwards, Mem. Bot. Surv. S. Afr.

36:274(1967).

H. aff. stenophylla sensu Killick, Mem. Bot. Surv. S. Afr.

34: 136 (1963); sensu Ross, l.c. 306 (1972).

Frutex ramosus, 40-150 cm altus: ramuli stellato-

tomentosi. Folia breviter petiolata; petiolus 1-2 mm

longus, dense stellato-tomentosus; lamina lanceolato-

elliptica, oblanceolato-elliptica vel lineari-elliptica,

7-20 mm longa, 2-7 mm lata, discolor, dense

stellato-tomentosa, supra grisea, subtus pallidior,

reticulata, apice obtuso vel rotundato, basi cuneata,

margine integra vel supra medium minute crenato-

dentato. Inflorescentia simplex vel basin versus parce

ramosa, 7-15 cm longa; rhachis stellato-tomentosa;

bracteae late ovatae, acutae, 4-7 mm longae, caducae,

stellato-pubescentes; verticillastri plerumque 6-flori;

pedicelli 2-3 mm longi, villosi. Calyx 5-7 mm longus,

villosus, glanduloso-punctatus; tubus 4-5 mm longus;

lobus posticus late ovatus, obtusus vel rotundatus,

suberectus, 2 mm longus, margine decurrente; dentes

laterales deltoideo-subulati, 1 mm longi; antici

lineari-subulati, 2 mm longi. Corolla pallide rosea vel

malvina, 8-11 mm longa, glabra vel extus labiis

puberulis; tubus 6-9 mm longus, rectus, apicem versus

sensim apliatus, ore 3 mm lato; labium posticum

parvum; anticum concavum, 3 mm longum. Stamina

9-12 mm exserta; postica prope basin tubi corollae

inserta, filamentis liberis prope basin minute pube-

scentibus, antica fauce corollae inserta, filamentis

fere ad apicem connatis. Stylus 10 mm exserta, apice

breviter bifidus.

Type. — Natal, 2829 (Harrismith), Cathedral Peak

Forest Research Station (-CC), Killick 1644 (PRE,

holo.).

Shrub, branched, 40-150 cm tall; branchlets

stellate-tomentose. Leaves shortly petiolate; petiole

1-2 mm long, densely stellate-tomentose; blade

lanceolate-elliptic to oblanceolate-elliptic or linear-

elliptic, 7-20 mm long, 2-7 mm broad, discolorous,

densely stellate-tomentose, dark grey above, paler

below, reticulate, apex obtuse to rounded, base

cuneate, margin entire or minutely crenate-dentate

above the middle. Inflorescence simple or sparingly

branched towards the base, 7-15 cm long; rhachis

stellate-tomentose; bracts broadly ovate, acute, 4-7

mm long, caducous, stellate-pubescent; verticillasters

usually 6-flowered; pedicels 2-3 mm long, villous.

Calyx 5-7 mm long, villous and freely gland-dotted;

tube 4-5 mm long; posticous lobe broadly ovate,

obtuse to rounded, suberect, 2 mm long, margin

decurrent; lateral teeth deltoid-subulate, 1 mm long;

anticous teeth linear-subulate, 2 mm long. Corolla

pinkish to mauve, 8-11 mm long, glabrous with

outer surfaces of lips puberulous; tube 6-9 mm long,

straight, widening gradually towards the apex, mouth

3 mm wide; posticous lip small; anticous lip concave,

3 mm long. Stamens exserted by 9-12 mm; posticous

stamens inserted near the base of the corolla tube,

filaments free, minutely pubescent near the base;

anticous stamens inserted at the throat of the corolla,

filaments united almost to the apex. Style exserted

by 10 mm, apex shortly bifid. Fig. 3.

Found at altitudes of 1 700 to 2 300 m in the

Natal Drakensberg between Mont-aux-Sources and

Cathkin Peak where it is a common shrub along

stream banks, at the foot of cliffs, and on mountain

sides. It flowers mainly from December to April.

Natal. — 2828 (Bethlehem): Royal Natal National Park

(-DB), Bayer & McClean 147; 217 ; Galpin 10168; Hutchinson,

Forbes & Verdoorn 52; Sidey 1655; Edwards 459; Trauseld 240;

1114. 2829 (Harrismith): Cathedral Peak (-CC), Killick 1644;

Hilliard & Burtt 3427. 2929 (Underberg): Cathkin Park (-AB),

Galpin 11846; above Champagne Hostel (-AB), Edwards 2300.

L. E. CODD

7

N \ FION A I.

I OK I A.

10. Heiltyzla -p. Klll’ck 1644 3 Oalpln 11S46, 10160 =

d=. it <U ■

\ *

at SSE' IIERUAR

rr^ ■/ Ia/jz.^ ,m

’-t

,»*oc<- 7-.

Fig. 3. — Hemizygia cinerea ( Killick 1644, PRE, holotype).

The few specimens of this species which Ashby

had at his disposal were confused with H. elliottii

(Bak.) Ashby and H. stenophylla (Guerke) Ashby,

and those specimens with entire leaf margins would

tend to run to H. elliottii in his key. However, H.

elliottii differs in having ovate-lanceolate leaves and a

capitate stigma, while the calyx is stellate-pubescent,

not villous as in H. cinerea. H. elliottii is essentially

a plant of hot, dry savanna country, extending from

Rhodesia to Botswana and to the western, northern

and eastern Transvaal lowveld, but does not enter

Natal.

H. stenophylla, on the other hand, has linear-

lanceolate leaves with somewhat thickened and

inrolled margins and colourful lanceolate bracts

12-15 mm long, while the calyx is glandular-hispid,

also lacking the villous hairs possessed by H. cinerea.

H. stenophylla occurs well to the south of H. cinerea

and at lower altitudes, from the Transkei to southern

Natal.

H. cinerea is probably more closely allied to the two

Transvaal species, H. rugosifolia Ashby and H.

incana Codd (described above). From H. rugosifolia

it differs in the dense tomentum on both surfaces of

the leaf, the smaller bracts and the villous calyx.

From H. incana it can be distinguished by the smaller

leaves, which are usually dark grey on the upper

surface and which are often entire or toothed only

in the upper half, while the flower parts (calyx,

corolla and stamens) are smaller. H. cinerea is known

only from the Natal Drakensberg between Cathkin

Park and Mont-aux-Sources, while H. incana

appears to be restricted to the Kaapsche Hoop area

in the eastern Transvaal.

7. Hemizygia elliottii (Bak.) Ashby in J. Bot.

Lond. 73: 345 (1935), pro parte excl. Natal spec.

Type: Matabeleland, Elliott s.n. (K, holo.).

Orthosiphon elliottii Bak. in FI. Trop. Afr. 5: 376 (1900).

O. messinensis Good in J. Bot. Lond. 63: 173 (1925). Type:

Transvaal, Messina, Moss & Rogers 153 (BM, holo.; PRE).

Soft shrub 35-60 cm tall, woody at the base;

branches stellate-tomentose. Leaves subsessile to

shortly petiolate; blade lanceolate to ovate, 15-25 mm

long, 4-12 mm broad, densely stellate grey velvety on

both surfaces; apex acute, base obtuse; margin entire.

Inflorescence simple or occasionally with a pair of

branches near the base, 6-12 cm long; rhachis densely

and shortly stellate-tomentose; bracts broadly ovate

to subrotund, persisting as a dense mauve-purple

coma, 7-11x5-8 mm, stellate-pubescent near the

truncate base, apex rounded; verticillasters 2-6-

flowered, up to 12 mm apart. Calyx 5 mm long,

sparingly stellate-tomentose mainly on the tube; upper

lip subrotund, rounded at the apex, 2 mm long,

markedly decurrent on the tube; lower 4 teeth deltoid-

subulate, up to 2 mm long. Corolla white to pale

mauve, 13 mm long, glabrous; tube 9 mm long,

widening to 3 mm at the throat; upper lip a small

appendage 1 mm long; lower lip concave, 4 mm long,

often deflexed. Stamens shortly exserted, not or only

slightly exceeding the lower corolla lip; upper pair

attached about the middle of the tube, filaments

pubescent for about two-thirds their length; lower

pair attached at the throat, adhering loosely at the

base for a few mm. Style capitate.

Found in dry, subtropical woodland in western,

northern and eastern Transvaal, at altitudes of 300

to 1 300 m, often on red sandy loam soil; also in

Botswana and Rhodesia. Collected in flower between

October and April.

Botswana. — 2326 (Mahalapye): Mahalapye (-BB), Yalala

143; Bayliss 1839 (NBG). 2425 (Gaberones): Pharing, Hillary

& Robertson 490; Miller B/975; 15 km S. of Gaberones (-DB),

Gillett 17480. 2426 (Mochudi): Mochudi (-AC), Harbor sub

Rogers 6554; Derdepoort (-CB), Codd 8857.

Transvaal. — 2229 (Waterpoort): Schroda (-AB), Pole

Evans 1963. 2230 (Messina) : Messina (-AC), Moss <Sc Rogers 153;

Wild 7632. 2329 (Petersburg) : between Bochem and Leipzig

(-AA), Bremekamp & Schweickerdt 150; near Bochem (-AC),

Bremekamp & Schweikerdt 46; 10 km N. E. of Pietersburg (-CD),

Codd 8324. 2330 (Tzaneen): near Blackhills (-DA), Oates 90.

2331 (Phalaborwa): 21 km N. of Letaba Camp (-DC), Codd «£

Dyer 4682. 2426 (Mochudi): 3 km S. of Rooibokkraal (-BB),

Leistner 3184; 22 km S.W. of Rooibokkraal (-BD), Codd 8658.

2427 (Thabazimbi) : near Sentrum (-AD), Vahrmeijer 1358;

farm Vaalpenskraal (-CB?), Theron & Marsh 251. 2428 (Nyl-

stroom): Towoomba Pasture Research Station (-CD), Irvine 56.

2430 (Pilgrim’s Rest): near Mica (-BB), Acocks 16754; Schlie-

ben 9348. 2431 (Acornhoek): between Klaserie and Acornhoek

(-CA), Rauh & Schlieben 9706; 13 km E. of Skukuza (-DC),

Codd 5036. 2526 (Zeerust): Zeerust (-CA), Pole Evans 407.

Superficially similar to H. cinerea Codd but the

small grey leaves are usually broader and the margins

are always entire. It is, however, not closely related

to H. cinerea because the stamen filaments are

pubescent to well above the middle (this can often

be seen on dried specimens without dissecting the

flowers), the stigma is capitate and there is a coma

of colourful bracts at the apex of the inflorescence.

An interesting variation is found in the numbers of

flowers produced per verticillaster. Normally a

constant character for a species, two series are found

in H. elliottii. In all specimens from Rhodesia, Bot-

swana, western and northern Transvaal, the verti-

cillasters are 2-flowered, while the specimens from the

8

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

eastern Transvaal lowveld have 6-flowered verti-

cillasters. As no other difference can be found it is

not proposed to accord separate rank for the latter

form.

8. Hemizygia gerrardii ( N.E . Br.) Ashby in J.

Bot. Lond. 73: 345 (1935); Ross, FI. Natal 306 (1972).

Type: Natal: “near Ingoma”, Gerrard 1239 (K, holo.).

Orthosiphon gerrardii N.E. Br. in FI. Cap. 5,1 : 249 (1910).

Soft branched shrub ca. 1 m. tall; branches stellate-

pubescent, glabrescent with age, bark flaking off in

strips. Leaves petiolate; petiole 1-3 mm long; blade

ovate to broadly elliptic, ca. 15x10 mm, thickish,

densely, and somewhat coarsely stellate grey velvety

on both surfaces; apex obtuse, base obtuse to truncate;

margin entire. Inflorescence usually simple, 4-5 cm

long; rhachis stellate-floccose; bracts broadly ellip-

tical, persisting as a mauve-purple coma, ca. 8 X 5 mm,

stellate-pubescent; verticillasters 2-flowered, 3-4 mm

apart. Calyx 8-10 mm long, stellate-tomentose;

upper lip ovate, 2 mm long, decurrent on the tube;

lower 4 teeth deltoid-subulate, the lowest pair dis-

tinctly the longest, 3 mm long. Corolla mauve-pink;

tube 17-20 mm long, pubescent; upper lip short;

lower lip concave, 6 mm long. Stamens well exserted;

upper pair attached near the throat; lower pair

attached at the throat, filaments connate to half-way

up.* Stigma entire or minutely bifid.

Known from two gatherings, one in northern Natal

and the other from southern Transvaal; found in

grass among rocks.

Transvaal. — 2731 (Louwsberg): 30 km S. of Sulphur Spring

(-AC), Dyer cfc Verdoorn 5829.

Natal. — Grid unknown: “near Ingoma”, Gerrard 1239 (K).

Resembles H. elliottii but the calyx and corolla

are distinctly larger.

9. Hemizygia floccosa Launert in Mitt. Bot.

Miinchen 7: 302 (1968); Launert & Schreiber in

Prodr. FI. S.W. Afr. 123: 13 (1969). Type: S.W.

Africa, Outjo, De Winter & Hardy 8139 (PRE,

holo.; M).

A soft shrublet 40-80 cm tall, woody below,

sparingly branched; branches pale reddish-brown,

loosely stellate-floccose, glabrescent with age. Leaves

petiolate; petiole 5-10 mm long; blade ovate 2,8-

4,5 cm long, 1,5-2, 2 cm broad, loosely to densely

stellate-floccose on both surfaces, nerves reticulate

below; apex subacute, base obtuse; margin obscurely

and somewhat distantly crenate-dentate. Inflorescence

simple or with a pair of branches near the base, lax;

rhachis glandular-puberulous; bracts deciduous, small,

ovate, 5x2,5 mm, pubescent; verticillasters 2-

flowered, 2-3 cm apart. Calyx 11 mm long, glandular-

strigose; upper lip broadly ovate, rounded, 2,5 mm

long, decurrent; lower 4 teeth deltoid subulate to

spinescent, the lowest pair the longest, 3 mm long.

Corolla pale mauve, 20 mm long, glabrous; tube

15 mm long widening from below the middle to 5 mm

wide at the throat; upper lip broad, 3 mm long;

lower lip concave, 5 mm long, usually deflexed.

Stamens shortly exserted, not exceeding the lower lip

of the corolla; upper pair attached near the middle

of the tube, scarcely exserted, filaments pubescent

below; lower pair attached at the throat, filaments

4 mm long, connate for about half their length.

Stigma capitate.

* Description of stamens taken largely from N. E. Brown, l.c.

A rare plant occurring along dry watercourses in

the central Namib area of South West Africa.

S.W. A. — 2014 (Welwitschia): near Bethanis (-AD), Giess

3929; 55 km W. of Welwitschia on road to Torra Bay (-BC),

De Winter d- Hardy 8139. 2114 (Uis): Brandberg (-AB), Lieben-

berg 5001.

Easily distinguished from other species in South

West Africa by the dense floccose pubescence of

stellate (branched) hairs on the relatively large

petiolate leaves, and 2-flowered verticillasters.

10. Hemizygia stenophylla ( Guerke ) Ashby in J.

Bot. Lond. 73: 347 (1935); Ross, FI. Natal, 306

(1972). Lectotype: Cape, E. Griqualand, near

Enyembe, Tyson in Herb. Austr. Afr. 1293 (K,

lecto.).

Orthosiphon stenophyllus Guerke in Bot. Jahrb. 26: 84

(1898); N.E. Br. in FI. Cap. 5,1 : 250 (1910).

Soft shrub 30-90 cm tall, branching from the

base; branches arising from a perennial woody

rootstock, ascending, sparingly to freely branched,

densely leafy, shortly stellate-tomentose. Leaves sub-

sessile; blade linear-lanceolate or elliptic-lanceolate

to lanceolate, 12-30 mm long, 3-5 mm broad, upper

surface, dark grey to blackish, finely and shortly

pubescent with nerves impressed, lower surface

densely grey stellate-velvety with nerves raised and

almost parallel to the main nerve: apex tapering

gradually, base obtuse; margin revolute, entire.

Inflorescence simple or with one or two pairs of

branches near the base, 8-18 cm long; rhachis

glandular-hispidulous often with some branched

hairs; bracts persisting as a colourful coma, lanceolate

to ovate-lanceolate, 10-15 mm long, acute, mauve

to purple, stellate-tomentose; verticillasters 4-6-

flowered. Calyx 7-8 mm long, glandular-hispidulous;

upper lip ovate, rounded, 3 mm long, decurrent;

lower 4 teeth subulate to bristle-like, the lowest pair

the longest, up to 4 mm long. Corolla pale mauve to

rosy-mauve, 13 mm long, glabrous except for the

lips; tube 10 mm long, widening to 3 mm at the

throat; upper lip a small appendage; lower lip

concave, 3 mm long, often deflexed. Stamens exserted

by 10 mm, well exceeding the lower lip; upper pair

attached about the middle of the tube, filaments

pubescent in the lower part; lower pair attached at the

throat, filaments united to or near the apex. Stigma

minutely bifid.

Found in dense grassland often near forest and

among rocks in southern Natal and East Griqualand.

Flowers mainly from January to March.

Natal. — 2930 (Pietermaritzburg): Boston (-CA), Medley

Wood 966 (SAM); 4624 (NH). 3029 (Kokstad): Mt. Ingeli,

near Weza (-DA), Medley Wood 3107 (NH); Strey 6300; 6334;

10923; Nicholson 1218.

Cape. — 3029 (Kokstad): Mt. Emyembe (-BA), Tyson in

Herb. Austr. -Afr. 1293 (SAM); 1720 (SAM); 1760; 2137 (SAM);

Insikeni Forest (-BA), Tonder 5; 16 km S. of Umzimkulu

(-BD), Codd 8568; between Brooks Nek and Bizana (-DA?),

Hilliard <fc Burtt 6748; near Fort Donald (-DC), Tyson 1666

(SAM).

Resembles H. rehmannii (Guerke) Ashby from the

north-eastern Transvaal but has slightly narrower,

more lanceolate leaves and a tuft of conspicuous

mauve-purple bracts at the apex of the inflorescence.

Although it has been confused with the species now

described as H. cinerea Codd, the latter tends to have

more elliptical leaves and also lacks the coma of

colourful bracts. Although both occur in Natal,

H. cinerea occurs further to the north and at higher

altitudes, between Cathedral Peak and Mont-aux-

Sources.

L. E. CODD

9

11. Hemizygia rehmannii ( Guerke ) Ashby in J.

Bot. Lond. 73: 347 (1935). Type: Transvaal, Hout-

boschberg, Rehmann 6172 (Z, holo; BM).

Orthosiphon rehmannii Guerke in Bull. Herb. Boiss. 6: 557

(1898); N.E. Br. in FI. Cap. 5,1: 251(1910).

Soft shrub branching from a perennial woody root-

stock, forming a round bush 30-80 cm tall; branches

erect or ascending, usually sparingly branched, with

dense short stellate hairs and villous hairs inter-

mingled, densely beset with leaves. Leaves sessile;

blade narrowly elliptic to oblong-elliptic, 10-22 mm

long, 3-8 mm broad, upper surface dark grey to

brownish, finely pubescent to rugose with nerves

impressed, lower surface densely grey to yellowish-

grey stellate-velvety with nerves raised; apex acute to

obtuse, base somewhat cuneate; margin revolute,

entire or finely crenate-dentate in the upper half.

Inflorescence simple or branched, 6-22 cm long;

rhachis finely to coarsely stellate-pubescent; bracts

small, ovate, acute, 5-6 mm long, stellate-pubescent;

verticillasters 6-flowered, 10 mm apart. Calyx 9-10 mm

long, glandular-hispid; upper lip ovate, rounded,

2,5 mm long, decurrent; lower 4 teeth weakly

subulate, the lowest pair the longest, 2,5 mm long.

Corolla pale mauve, 17 mm long, glabrous except for

the lips; tube 14 mm long, widening to 4 mm at the

throat; upper lip a small appendage; lower lip con-

cave, 3 mm long, eventually deflexed. Stamens exserted

by 10 mm, well exceeding the lower lip; upper pair

attached below the middle of the tube, filaments

glabrous; lower pair attached at the throat, filaments

united to near the apex. Stigma minutely bifid.

Occurs usually in shallow sandy soil among rocks

in grassland, often near forest margins, from Wood-

bush to The Downs in north-eastern Transvaal, at

altitudes of 1 500-2 000 m. Flowering is mainly from

January to March.

Transvaal. — 2329 (Pietersburg): Houtboschberg (-DD)’

Schlechter 4442; Iron Crown Mt. (-DD), Mogg 16652; Wolk-

berg (-DD), Meeuse 9866; near Houtboschdorp (-DD),

Codd 9426. 2330 (Tzaneen): Westfalia Estate (-CA), Scheepers

909; Woodbush (-CC), Pole Evans 4746; Pott sub TRV 13393;

New Agatha (-CC), McCallum s.n.; Muller d- Scheepers 111.

2429 (Zebediela): Ashmoledales (-BB), Pole Evans H. 19010.

2430 (Pilgrim’s Rest): mountains near Trichardsdal and The

Downs (-AA), Vahrmeijer 2369.

Its affinity with H. stenophylla is discussed under

that species. From H. cinerea, a Natal species which

also has small, inconspicuous bracts, it differs in the

leaves being rather more rugose above and usually

greyish-yellow below, and the larger corolla, which is

only 8-11 mm long in H. cinerea as against about

17 mm long in H. rehmannii.

12. Hemizygia subvelutina ( Guerke ) Ashby in J.

Bot. Lond. 73: 346 (1935). Type: Lydenburg, near

Paarde Plaats, Wilms 1152 (BM, K).

Orthosiphon subvelutinus Guerke in Bot. Jahrb. 26: 80

(1898); N.E. Br. in FI. Cap 5,1: 253 (1910). O. heterophyllus

Guerke, l.c. 82 (1898). Syntypes: eastern Transvaal, near

Spitzkop, Wilms 1 148 (BM); 1155 (K).

Bushy herb or soft shrublet 20-50 (-80) cm tall,

branching from the base; branches few to many from

a perennial woody rootstock, erect or ascending,

sparingly branched, densely beset with leaves and

short leafy shoots, densely stellate-pubescent, often

with a yellowish tinge. Leaves sessile, usually ericoid,

linear to linear-lanceolate, 5-10 (-15) mm long, 1-2

(-5) mm broad, coriaceous, stellate-scabrid above,

usually yellowish stellate-tomentose below; margin

revolute, entire. Inflorescence simple, 5-11 cm long;

rhachis densely stellate-hispid with branched, usually

yellowish hairs; bracts persistent, 4-7x2-3 mm,

stellate-hispid, somewhat leaf-like; verticillasters 4-6-

flowered, occasionally less, 5-10 mm apart. Calyx

5-6 mm long, stellate-hispid; upper lip ovate, acute,

2 mm long, decurrent; lower 4 teeth deltoid-subulate,

up to 2 mm long. Corolla white, often tinged with

mauve, 12-16 mm long; tube 10-12 mm long, tubular,

2.5 mm wide, often slightly constricted at the throat,

sparingly pubescent (densely so on the lips); upper lip

1.5 mm long; lower lip shallowly concave, 2-4 mm

long. Stamens shortly exserted by 1,5-3 mm, not or

scarcely exceeding the lower lip of the corolla; upper

pair attached below the middle of the tube, filaments

puberulous near the base; lower pair attached at the

throat, filaments united only near the base or to

about half their length. Stigma shortly bifid.

Localized on the eastern Transvaal mountains

from Lydenburg and Pilgrim’s Rest to Kaapsche

Hoop, in dense grass among quartzite rocks and in

rock crevices, at altitudes of 1 400 to 2 200 m. Flowers

from November to March.

Transvaal. — 2430 (Pilgrim’s Rest): Pilgrim’s Rest (-DD),

Rogers 14321; 14871; 18328; Galpin 14447; Graskop (-DD),

Pole Evans 128; Holt 366; between Kowyns Pass and Sabie

(-DD?), Gillett 1017; Mac Mac Falls (-DD), Burn Davy 2536;

Codd 6446; 9480. 2530 (Lydenburg): farm Zwagershoek,

S.W. of Lydenburg (-AB), Obermeyer 327; Mount Anderson

(- BA), Smuts d- Gillett 2403; Meeuse 10074; Long Tom Pass,

Werdermann d- Oberdieck 2103; Leistner d- Mauve 3224; Kemps

Heights, 24 km S.W. of Lydenburg (-BA), Codd 8306; Sabie

Valley (-BA), Galpin 13811; Sabie (-BB), Rogers sub TRV

14869; Wager A118; Witklip (-BD), Kluge 301; 27 km S.E. of

Machadodorp (-CB), Bruce 487; Ohrigstad Nature Reserve

(-DC), Jacobsen 1493; Kaapsche Hoop (-DB), Pole Evans 985;

Wager sub TRV 15561; Codd 5751.

Closely related to H. teucriifolia (Hochst.) Briq. as

will be seen from the tubular corolla which is slightly

constricted at the mouth, and the very shortly exserted

stamens. It differs from H. teucriifolia in a few minor

respects, namely, the narrower, “ericoid” leaves

(although occasional broader, ovate leaves may be

present), the shorter internodes, the tendency for the

tomentum to be yellower in colour and the usually

pubescent corolla tube. The two meet in the Transvaal,

but do not appear to overlap; H. subvelutina is

restricted to the mountains from near Lydenburg and

Pilgrim’s Rest to Kaapsche Hoop, while H. teucriifolia

is distributed from the eastern Cape Province through

Natal to the Barberton area, appearing again in the

mountains of eastern Rhodesia.

There is some indication of introgression in the

south-eastern Transvaal. Specimens from Kaapsche

Hoop tend to have longer internodes and, occasionally,

glabrous corolla tubes, while an occasional specimen

from near Barberton may have pubescence on the

corolla tube (apart from ihe lips which are always

pubescent on the outer surface). Usually leaf shape and

the colour of the pubescence can be used as a guide

in such cases and it is felt that both species can

justifiably be upheld. See also notes under H. albiflora

(p. 10).

N. E. Brown, l.c., considered that in some

specimens the filaments of the anterior pair of stamens

were free to the base. This was not confirmed in the

present study though admitedly more flowers should

be examined than is possible in a herbarium. The

degree of union was variable, from about 0,5 mm to

1 ,5 mm, the latter representing about half the length

of the filaments.

13. Hemizygia teucriifolia (Hochst.) Briq. in

Pflanzenfam. 4, 3a: 369 (1897); Annu. Conserv.

Jard. Bot. Geneve 2: 247 (1898); Ashby in J. Bot.

Lond. 73: 346 (1935); Ross, FI. Natal 306 (1972).

Type: Natal, Table Mt., Krauss 448 (BM, K).

10

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

Ocimum teucriifolia Hochst. in Flora 28: 66 (1845); Benth.

in DC., Prodr. 12: 41 (1848).

Orthosiphon woodii Guerke in Bot. Jahrb. 26: 83 (1898). Type:

Natal, Entumeni, Medley Wood sub NH 783 (SAM) (= Medley

Wood 3964 in K, NH). O. galpiniana Briq. in Bull. Herb. Boiss.

ser. 2,3: 993 (1903). Type: Transvaal, Barberton, Saddleback,

Galpin 1217 (K, NH, PRE). O. teucriifolius (Hochst.) N.E.

Br. in FI. Cap. 5,1: 254 (1910). — var. galpinianus (Briq.) N.E.

Br., l.c. 254(1910).

Bushy herb 15-30 cm tall, branching freely from a

woody, perennial base; branches erect or ascending,

usually simple, greyish stellate-pubescent. Leaves

linear to lanceolate or elliptic, 8-18 mm long, 3-6 mm

broad, stiffish, stellate-scabrid and blackish above,

greyish stellate-tomentose below; apex acute, base

obtuse; margin revolute, entire. Inflorescence simple,

4-8 cm long, fairly lax or congested; rhachis stellate-

villous; bracts persistent, 4-6 x 2-3 mm, stellate-

hispid, somewhat leaf-like; verticillasters 4— 6-flowered,

2-10 mm apart. Calyx 5-6 mm long, stellate-pube-

scent; upper lip ovate, rounded, 2 mm long, decurrent;

lower 4 teeth deltoid-subulate, up to 2 mm long.

Corolla mauve, 10-12 mm long; tube 9-10 mm long,

tubular, 2,5 mm wide, slightly constricted at the

throat, glabrous (pubescent on the lips); upper lip

1,5 mm long; lower lip slightly concave 2 mm long.

Stamens shortly exserted by 1-3 mm or the upper pair

scarcely exserted; upper pair attached below the

middle of the tube, filaments puberulous in the lower

half; lower pair attached at the throat, filaments

united for half or more of their length. Stigma

minutely bifid.

Locally frequent in mountain grassland at altitudes

of 600 to 1 500 m at scattered localities from Stutter-

heim District in the eastern Cape, to Natal and the

Barberton District of the Transvaal, appearing again

in the mountains of eastern Rhodesia.

Transvaal. — 2530 (Lydenburg): Nelshoogte (-DB), Midler

2151 ; Barker s.n. (NBG). 2531 (Komatipoort): Saddleback Mt.

(-CC), Galpin 1217; Lomati Valley (-DA), Thorncroft 2013.

Natal. — 2731 (Louwsburg): Ngome (-CD), Gertsner 5120’

Strey 9338. 2831 (Nkandla): Babanango (-AC), King 296’

Acocks 11781; Codd 1773; Nkandla (-CA), Wylie sub NH 9405

(NH); Melmoth (-CB), Mogg 6059; Ntumeni (-CD), Medley

Wood sub NH 783 (SAM); 3964 (NH). 2930 (Pietermaritzburg):

near Howick (-AC), Medley Wood 8730; near The Dargle

(-AC), Hilliard & Burtt 3193; Swartkop (-CB), Wylie sub Wood

10106; Fisher 703; near Hela Hela (-CC), Strey 9217; near

Richmond (-CD), Medley Wood 10068. 3029 (Kokstad):

Ngeli Mt. (-DA), Hilliard <£■ Burtt 3483. 3030 (Port Shepstone):

Campbellton (-AD), Rudatis 1790; Illovo (-BB?), Medley

Wood 1877 (NH).

Cape. — 3029 (Kokstad): Zuurberg (-BA), Tyson 1561.

3227 (Stutterheim): Mt. Kemp (-CB), Sim s.n.; 19595; Pirie

(-CC), Sim 107.

Its relationship with H. subvelutina is discussed

under that species. It also resembles the following

species, H. albiflora, which can readily be separated by

its lack of stellate (branched) hairs.

Apparently H. teucriifolia is not readily eaten by

stock and tends to increase on veld which is subjected

to overgrazing.

14. Hemizygia albiflora (N.E. Br.) Ashby in J.

Bot. Lond. 73: 348 (1935): Type: Transvaal, Mac

Mac, Mucld s.n. (K., holo.).

Ortosiphon albiflorus N.E. Br., in FI. Cap. 5,1: 251 (1910).

O. decipiens N.E. Br., I.c. 252 (1910). Type: Transvaal, Mac

Mac, Mudd s.n. (K,holo.).

Hemizygia rehmannii sensu Compton, FI. Swaz. 67 (1966).

Woody shrublet 30-150 cm tall; branches markedly

woody towards the base, somewhat gnarled and

branching, procumbent to ascending, villous, densely

beset with leaves and short leafy shoots. Leaves sessile,

ericoid; blade linear to linear-elliptic, 10-30 mm

long, 1,5-5 mm broad, stiff, subglabrous and shiny

to appressed villous and blackish above, densely

appressed villous below with long white matted hairs;

apex and base tapering; margin strongly revolute,

entire. Inflorescence simple or with a pair of branches

near the base, 5-10 cm long; rhachis glandular-

hispid; bracts deciduous, ovate, acute, 5-8 x 3-5 mm,

sparingly pubescent; verticillasters mainly 6-flowered,

5-15 mm apart. Calyx 6-8 mm long, glandular-hispid;

upper lip ovate, rounded, 2 mm long, decurrent;

lower 4 teeth deltoid-subulate, up to 2 mm long.

Corolla white, 12-15 mm long; tube 10-12 mm long,

curved, glabrous, tubular, expanding to 3 mm wide

about the middle and then parallel sided or slightly

constricted at the mouth, upper lip very small, less

than 1 mm long; lower lip concave, 2-3 mm long

Stamens exserted by 4-6 mm, exceeding the lower lip

of the corolla; upper pair attached below the middle

of the tube, filaments finely puberulous in the lower

half; upper pair attached at the throat, filaments

united for almost their entire length. Stigma minutely

bifid.

Found among quartzite rocks, often with semi-

alpine flora at altitudes of 1 800 to 2 400 m, in the

mountains of the eastern Transvaal and extending to

northern Swaziland. The main flowering season is

from November to March.

Transvaal. — 2430 (Pilgrim’s Rest): Mariepskop (-DB)

Smuts <L GUlett 3598; Codd 7853; Meeuse 9960; Van der Schijff

4366; 4777; 4827; 4922; 6189; 6340; Werdermann dk Oberdieck

1838; 1905; Killick dk Strey 2390; 2400; Blyde River Gorge

(-DB), Galpin 14628; Hebronberg (-DB), Hilliard dk Burtt 6002:

God’s Window, N. of Pilgrim’s Rest (-DD), Davidson dk Mogg

32922; 33473; Graskop (-DD), Burtt Davy 1478; Galpin 14352;

between Graskop and Mac Mac (-DD), Loock s.n.; Mac Mac

(-DD), Nel 262. 2530 (Lydenburg): Steenkampsberg, 46 km W.

of Lydenburg (-AA), Codd 8207; 13 km E. of Draaikraal (-AA),

Bruce 328; Dullstroom, Suikerboskop (-AC), Galpin 13052;

13068; Mt. Anderson (-BA), Pole Evans 4296 (59); Smuts 4k

Gillett 2419; 2430; Long Tom Pass (-BA), Werdermann dk

Oberdieck 2088; Bakenkop 14 km E. of Sabie (-BB), Leistner &

Mauve 3235; Schoemanskloof, Rietvlei (-BC), Smuts 5; 13 km

N.W. of Belfast (-CA), Story 5345; Mareskop, near Machado-

dorp (-CA), Bruce 518; near Waterval Boven (-CB), Codd

10481; 25 km S.E. of Machadodorp (-CB), Codd 8269; 8270;

Kaapschehoop (-DB), Pole Evans 983; Rogers 19571 ; Thode A

1636. 2531 (Komatipoort): between Barberton and Havelock

(-CC), Hilliard dk Burtt 3663. 2630 (Carolina): Ermelo, Billy’s

Vlei (-CA), Burtt Davy 8092.

Swaziland. — 2631 (Mbabane): Ngwenya Mts. (-AC)

Compton 26683.

The hard, narrow (“ericoid”) leaves superficially,

resemble those of H. subvelutina and H. teucriifolia

but are easily distinguished by the lack of stellate

(branched) hairs. H. albiflora is a woody, freely

branched plant which grows among rocks where it is

fairly well protected against fire, whereas the other

two species form annually a number of slender stems

from a thick woody rootstock in response to repeated

grass fires. LI. albiflora also has a slightly longer corolla

and the stamens are exserted beyond the lower corolla

lip, while the filaments of the lower pair of stamens

are united nearly to the apex.

15. Hemizygia pretoriae ( Guerke ) Ashby in J.

Bot. Lond. 73: 356 (1935). Type: Transvaal, Pretoria,

Wilms 1151 (BM).

Bushy herb 10-30 cm tall, branching freely from a

perennial woody base; branches arising annually,

several to many, simple, erect or ascending, hispid to

villous, sometimes with branched hairs intermingled

(subsp. heterotricha). Leaves subsessile to shortly

petiolate; blade narrowly elliptic or oblanceolate to

ovate, or subrotund, 8-24 mm long, 2-15 mm broad,

subglabrous to villous, sometimes with stellate

(branched) hairs, conspicuously gland-dotted, often

folding along the midrib; apex acute to obtuse, base

cuneate, often tapering gradually; margin entire or

rarely with a few small teeth. Inflorescence simple

L. E. CODD

11

4-8 cm long, medium-dense; rhachis glandular-

hispid to villous; bracts persistent, leaflike, ca. 10x4

mm, hispid to stellate-pubescent; verticillasters (2-)

4-6-flowered, 10-20 mm apart. Calyx 7-8 mm long at

flowering, enlarging considerably in fruit, glandular-

hispid; upper lip ovate-elliptic, rounded, 2,5 mm long,

decurrent; lower 4 teeth deltoid-subulate, up to

2,5 mm long, the lower pair in particular becoming

bristle-like. Corolla whitish to pale mauve, 14-16 mm

long, puberulous; tube 10-12 mm long, narrowly

tubular, scarcely widening to 2 mm wide at the

throat; upper lip relatively long and narrow, 3 mm

long; lower lip shallowly concave, 4 mm long.

Stamens with only the lower pair shortly exserted by

2-3 mm (less than the lower lip); upper pair included,

attached near or above the middle of the tube,

filaments very slender, glabrous; lower stamens

attached at the throat, united for more than half

their length. Stigma minutely bilobed.

Distributed from central to eastern Transvaab

Swaziland and northern Natal, in dense grassland,

often among rocks, mainly at altitudes of 1 000 to

1 800 m. The species is characterized by the narrowly

tubular corolla, scarcely expanding towards the

throat; the longish upper lip of the corolla which

almost equals the lower lip; and the fact that the

upper pair of stamens are never exserted. In certain

other species the upper pair of stamens are very short

but eventually they are slightly exserted, e.g. H.

subvelutina, H. teucriifolia and H. persimilis.

Two subspecies are recognized which are separated

mainly on the presence or absence of stellate

(branched) hairs (see key to species, p. 2) and to

some extent of leaf shape.

(a) subsp. pretoriae.

Hemizygia pretoriae (Guerke) Ashby in J. Bot. Lond. 73:

356 (1935); Compton, FI. Swaz. 67 (1966), pro parte; Ross,

FI. Natal 306 (1972). Type: Transvaal, Pretoria, Wilms 1151

(BM).

Orthosiphon pretoriae Guerke in Bot. Jahrb. 26: 81 (1898);

N.E. Br. in FI. Cap. 5,1: 254 (1910). O. natalensis Guerke,

l.c. 82 (1898). Syntypes: Natal, Glencoe, Medley Wood 4756

(K, NH); Kuntze s.n.; Coldstream, Rehmann 6918.

Stellate (branched) hairs absent throughout the

plant; leaves narrowly elliptic to oblanceolate or

obovate.

Distribution and ecology as for the species, though

absent from southern Swaziland; flowering is mainly

from October to February.

Transvaal. — 2527 (Rustenburg): Uitkomst (-DC), Coetzee

416; Jack Scott Nature Reserve (-DD), Wells 2394. 2528

(Pretoria): near Silverton (-CB), Phillips 3028; Baviaanspoort

(-CB), C. A. Smith 1062: Rietvlei Dam area (-CD), Pole Evans

287; Acocks 11256; Codd 2195; Meeuse 9249; Tygerpoort

(-CD), Repton 608; Premier Mine (-DA), Menzies s.n.; Rayton

(-DA), Rogers 21806; Bronkhorstspruit (-DC), Janse 59.

2529 (Witbank): Loskopdam (-AD), Theron869; near Monster-

lus (-BA), Acocks 20880; near Witbank, Gilfillan sub. Galpin

7233; Bruce 95; near Middelburg (-CD), Jenkins sub TRY

10700; Hewitt sub TRY 10431; Young A23; Codd 10079;

Doornkop 273 (-CD), Du Plessis 616. 2530 (Lydenburg): near

Helvetia North (-CB), Young A261 ; near Machadodorp (-CB),

Jenkins sub TRY 12705; 15 km W. of Machadodorp (-CB),

Codd 8027. 2627 (Potchefstroom): Little Falls (-BB), Mogg

20249 (J). 2628 (Johannesburg): Johannesburg (-AA), Gilfillan

sub Galpin 1440; Leendertz 1721 ; Bryant D52; Gertner 6720;

Macnae 1244 (J); Lucas 70 (J); Moss 2803 (J); 2804 (J )\2389 (J);

2770 (J); Kaalfontein (-AB), Pole Evans H 13532; H 16794.

2629 (Bethal): near Ermelo (-DB), Pott sub TRY 15112. 2630

(Carolina): Bushmans River Valley (-AA), Galpin 12442.

2730 (Vryheid): Mooihoek (-AB), Devenish 1052; Piet Retief

(-BB), Galpin 9645.

Swaziland. — 2631 (Mbabane): Mbabane (-AC), Compton

26785.

Natal. — 2729 (Volksrust): Newcastle (-DD), Schlechter

3420. 2730 (Vryheid): “Retirement” (-AD?), Devenish 1335;

near Grootspruit (-BC), Strey 8042; Kaffir Drift (-CD),

Thode A244. 2830 (Dundee): near Glencoe (-AA), Medley

Wood 4756; Dundee (-AA), Edwards 1084. 2831 (Nkandlap

20 km E. of Babanango (-AC), Codd 1775.

(b) subsp. heterotricha Codd, var. nov. a typica

foliis stellato-pubescentibus differt.

Type: Swaziland, 2631 (Mbabane), near Hlatikulu

(-CD), Compton 26320 (PRE, holo.).

Stellate (branched) hairs present on stems, leaves

and bracts; leaves ovate to ovate-rotund; florally in

no way different from the typical. Fig. 4.

>A

Fig. 4. — Hemizygia pretoriae var. heterotricha ( Compton 26320,

PRE, holotype).

Found in south-western Swaziland, the Piet Retief

District of Transvaal and the Hluhluwe area of Natal;

recorded in flower from October to January.

Transvaal. — 2731 (Lowsburg): 32 km W. of Pongola

Settlement on road to Piet Retief (-AD), Acocks 13154; Codd

2102.

Swaziland. — 2631 (Mbabane): near Mankaiana. (-CA),

Compton 30458; near Hlatikulu (-CD), Compton 26259; 26320;

28323; 29251.

Natal. — 2832 (Mtubatuba): Hluhluwe Game Reserve

(-AA), Ward 3960.

Normally, subsp. heterotricha does not differ in

floral characters or growth habit from the typical

and thus subspecific status is considered appropriate.

It occurs slightly to the south-west of subsp. pretoriae

and no overlapping in distribution has yet been found.

The specimen Ward 3960 has an unusual habit

consisting of a slender branched stem about 30 cm

tall, apparently not arising from a woody rootstock

12

THE SOUTH AFRICAN SPECIES OF HEM1ZYGIA (LAMIACEAE)

as in the normal behaviour of the species. Florally it

does not deviate from H. pretoriae and it possesses

the stellate pubescence of subsp. heterotricha. The

unusual habit may be the result of the absence of

fire over a period of years or it may represent a distinct

ecological form adapted to the lower altitude.

16. Hemizygia modesta Codd, sp. nov. a H.

thorncroftii (N.E. Br.) Ashby foliis brevioribus,

bracteis parvioribus, inconspicuis, verticillastris 2-

floribus differt.

Fruticulus 12-25 cm altus; caules annui, graciles,

erecti, caudice lignoso exoriens, hispidi vel villosi.

Folia sessilia vel subsessilia, late ovata, ovata, elliptica

vel lanceolato-elliptica, 6-12 mm longa, 4-6 mm lata,

parce vel dense hispida, glanduloso-punctata, nervis

obscuris, apice acuto vel obtuso, basi obtusa, margine

integro. Inflorescentia simplex, 5-10 cm longa;

rhachis glanduloso-hispidu ; vel villosus; bracteae

ovatae, acuminatae, 4-5 mm longae, inconspicuae,

cuducae, hispidae vel villosae, glanduloso-punctatae;

verticillastri 2-flori; pedicelli 2 mm longi. Calyx per

anthesin 7-8 mm longus, hispidus, copiose glanduloso-

punctatus; tubus 4-5 mm longus; lobus posticus late

ovatus, suberectus, obtusus vel rotundatus, 2 mm

longus, margine decurrenti; dentes laterales deltoideo-

subulati, 2 mm longi; antici linear-subulati, 2,5-3 mm

longi. Corolla alba vel pallido-malvina, 15-16 mm

longa, glabra vel extus labiis puberulis; tubus 11-

12 mm longus, apicem versus sensim ampliatus, ore

4 mm lato; labium posticum 1,5 mm longum;

anticum concavum, 4-5 mm longum, horizontale vel

recurvum. Stamina 10 mm exserta; postica circa

medium tubi corollae inserta, filamentis libris basin

versus pubescentibus; antica fauce corollae inserta,

filamentis ad apicem connatis. Stylus 15 mm exsertus,

apice breviter bifido.

Type. — Swaziland, 2631 (Mbabane), Bomvu Ridge

(-AA), Compton 28368 (PRE, holo.).

Shrublet 12-25 cm tall; stems annual, slender,

erect, arising from a woody rootstock, hispid to

villous. Leaves sessile or subsessile, broadly ovate to

elliptic or lanceolate-elliptic, 6-12 mm long, 4-6 mm

broad, sparingly to densely hispid, gland-dotted,

nerves indistinct, apex acute to obtuse, base obtuse,

margin entire. Inflorescence simple, 5-10 cm long;

rhachis glandular-hispid to villous; bracts ovate,

acuminate, 4-5 mm long, inconspicuous, caducous,

hispid to villous, gland-dotted; verticillasters 2-

flowered; pedicels 2 mm long. Calyx when flowering

7-8 mm long, hispid, freely gland-dotted; tube

4-5 mm long; posticous lobe broadly ovate, suberect,

obtuse to rounded, 2 mm long, margin decurrent;

lateral teeth deltoid-subulate, 2 mm long; anticous

teeth linear-subulate, 2,5-3 mm long. Corolla white

to pale mauve, glabrous with outer surfaces of lips

puberulous; tube 11-12 mm long, widening gradually

towards the apex, mouth 4 mm wide; posticous lip

1,5 mm long; anticous lip concave, 4-5 mm long,

horizontal to recurved. Stamens exserted by 10 mm;

posticous stamens inserted about the middle of the

corolla tube, filaments free, pubescent towards the

base; anticous stamens inserted in the throat of the

corolla, filaments united to the apex. Style exserted

by 15 mm, apex shortly bifid. Fig. 5.

Found in mountain grassland subjected to periodic

burning, in the mountains behind Barberton, in

Swaziland and in the Piet Retief district; flowering

takes place in spring while the grass is still short,

though it can continue until later depending on local

conditions.

Fig. 5. — Hemizygia modesta ( Compton 28368, PRE, holotype).

Transvaal. — 2531 (Komatipoort): Cythna Letty Reserve

(-CC), Mauve 4807; 16 km S.E. of Barberton on road to Have-

lock (-CC), Acocks 12867; Codd 1623. 2730 (Vryheid): Piet

Retief (-BB), Leipoldt s.n.

Swaziland. — 2531 (Komatipoort): Havelock (-CC), Comp-

ton 29123. 2631 (Mbabane): Bomvu Ridge (-AA), Compton

28368; Forbes Reef (-AC), Compton 30975; 32443; Mankaiana,

near Gege (-CA), Compton 30013.

In habit and ecology H. modesta resembles H.

pretoriae (Guerke) Ashby and H. thorncroftii (N.E.

Br.) Ashby in being small, spring-flowering shrublets,

sending up annual shoots from a woody rootstock

often before the first rains occur, and is most

noticeable after the grass has been burnt.

From H. pretoriae it is readily distinguished on the

floral characters: in H. pretoriae the verticillasters are

6-flowered, the corolla scarcely widens towards the

mouth and the upper pair of stamens remains included

in the corolla tube; in H. modesta the verticillasters

are 2-flowered, while the corolla widens towards the

mouth and all four stamens are well exserted.

The individual flowers of H. thorncroftii are similar

to those of H. modesta but again the verticillasters

are 6-flowered, the leaves are long and narrow

(1,5-3, 5 cm long), and the bracts are longer (8-

20 mm) and more colourful, persisting as a con-

spicuous apical coma. An occasional specimen of

H. thorncroftii has leaves shorter and wider than

usual and such specimens appear to be somewhat

intermediate, but the 6-flowered verticillasters and

conspicuous bracts place them without doubt in

H. thorncroftii.

L. E. CODD

13

Another dwarf species of the eastern Transvaal

highveld, S. foloisa S. Moore, has 2-flowered verti-

cillasters, but the leaves are much larger (2-6 x

1-3 cm), the inflorescence is usually paniculate and

the bracts are large and conspicuous.

There is a certain amount of variation included in

the specimens of H. modesta cited above, which calls

for further investigation. Three specimens from near

Havelock, Acocks 12867, Codd 1623 and Compton

29123, have shorter and broader leaves with more

villous pubescence on stems and leaves than the

typical form. However, Mauve 4807 from near

Barberton is somewhat intermediate and so separate

rank for the villous form does not seem to be justified.

The specimen Leipoldt s.n. from Piet Retief is more

robust than usual for the species. It was given the

manuscript name H. nervosa by Ashby, who refrained

from publishing it until further material became

available.

17. Hemizygia punctata Codd, sp. nov. a H.

transvaalensis (Schltr.) Ashby foliis, bracteis et

floribus parvioribus, verticillastris 2-floribus differt

Frutex ramosus, 60-120 cm altus; ramuli graciles,

hispiduli. Folia subcoriacea, breviter petiolata;

petiolus 1-2 mm longus; lamina elliptica vel elliptico-

oblanceolata, 10-15 mm longa, 3-7 mm lata, utrinque

hispidula, glanduloso-punctata, nervis obscuris, apice

acuto vel obtuso, basi cuneata, margine integro vel

supra medium minute crenato-dentato. Inflorescentia

simplex vel basin versus parce ramosa, 8-15 cm

longa; rhachis breviter hispidus; bracteae late ovatae,

4-6 mm longae, caducae, parce pubescentes; verti-

cillastri 2-flori; pedicelli 3 mm longi, hispidi. Calyx

per anthesin 6-8 mm longus, glanduloso-hispidus;

tubus 4 mm longus; lobus posticus late ovatus,

obtusus, suberectus, 2 mm longus, margine decurrenti;

dentes laterales deltoideo-subulati, 1,5 mm longi;

antici subulati, 2,5 mm longi. Corolla pallide malvina,

9-12 mm longa, glabra; tubus 7-10 mm longus,

rectus, apicem versus sensim ampliatus, ore 4 mm lato;

labium posticum parvum; anticum concavum, 2 mm

longum. Stamina 7-9 mm exserta; postica prope basin

tubi corollae inserta, filamentis libris, basin versus

pubescentibus ; antica fauce corollae inserta, filamentis

ad apicem connatis. Stylus 7-8 mm exsertus, apice

breviter bilobato.

Type: Transvaal, 2530 (Lydenburg), 18 km S.W-

of Lydenburg (-AB), Codd 8038 (PRE, holo.).

Shrub, branching, 60-120 cm tall; branchlets

slender, hispidulous. Leaves subcoriaceous, shortly

petiolate; petiole 1-2 mm long; blade elliptic to

elliptic-oblanceolate, 10-15 mm long, 3-7 mm broad,

both surfaces hispidulous and gland-dotted with the

nerves obscure, apex acute to obtuse, base cuneate,

margin entire or minutely crenate dentate above the

middle. Inflorescence simple or sparingly branched

towards the base, 8-15 cm long; rhachis shortly

hispid; bracts broadly ovate, 4-6 mm long, caducous,

sparingly pubescent; verticillasters 2-flowered; pedicels

3 mm long, hispid. Calyx when flowering 6-8 mm

long, glandular-hispid; tube 4 mm long; posticous

lobe broadly ovate, obtuse, suberect, 2 mm long,

margin decurrent; lateral teeth deltoid-subulate,

1,5 mm long; anticous teeth subulate, 2,5 mm

long. Corolla pale mauve, 9-12 mm long, glabrous;

tube 7-10 mm long, straight, widening gradually

towards the apex, mouth 4 mm wide; posticous lip

small; anticous lip concave, 2 mm long. Stamens

exserted by 7-9 mm; posticous stamens inserted near

the base of the corolla tube, filaments free, pubescent

towards the base; anticous stamens inserted in the

throat of the corolla, filaments united to the ape*.

Style exserted by 7-8 mm, apex shortly bilobed.

Fig. 6.

NATION A/, HHOSAkllJM.

nWTltlCT /

Fig. 6. — Hemizygia punctata ( Codd 8038, PRE, holotype).

Grows on stony slopes in grassland, often with

scattered trees and shrubs, and has been recorded

from the Lydenburg, Nelspruit and Barberton

Districts.

Transvaal. — 2530 (Lydenburg): 18 km S.W. of Lydenburg

(-AB), Codd 8038; Amajuba Mt., Schagen (-BD), Liebenberg

3088; above Rivulets Station, (-BD), Liebenberg 3323. 2531

(Komatipoort): Thorncroft Nature Reserve, off Agnes Mine

road (-CC), Edwards 4113.

Although superficially resembling the small-leaved

form of H. transvaalensis (Schltr.) Ashby (described

as Ocimum wilmsii Guerke, but now included in

H. transvaalensis), H. punctata differs in many

details. It is a slender-stemmed shrub, branching

above, in contrast to H. transvaalensis which branches

mainly from the base; the leaves are smaller, subentire

to minutely toothed in the upper half, the nerves are

indistinct and minute gland-dots are impressed in

both the upper and lower surfaces; the flowers are

smaller; and the bracts are small and inconspicuous

Ashby attached the manuscript name H. glan-

dulifolia to the specimen Liebenberg 3323, which he

saw in 1936, but refrained from describing due to

inadequate material.

14

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

18. Hemizygia bolusii (N.E. Br.) Codd in Bothalia

8: 159 (1964); Ross, FI. Natal 306 (1972). Type:

Natal, Giants Castle, A. Bolus in Herb. Guthrie

4894 (BOL!, holo.).

Orthosiphon bolusii N.E.Br. in FI. Cap. 5,1 : 258 (1910).

Stems several, erect, 25-30 cm tall arising annually

from a woody perennial base, softly woody at the

base, herbaceous above, sparingly branched, villous.

Leaves petiolate; petiole 2-4 mm long; blade ovate,

2-2,5 cm long, 1,4- 1,8 cm broad, brownish and

appressed hispid above, paler, hispid to villous

especially on the nerves and minutely gland-dotted

below; apex and base obtuse to rounded; margin

with a few minute teeth above the middle. Inflorescence

simple, 10-14 cm long, lax; rhachis glandular hispid;

bracts caducous, inconspicuous, ovate, 4-5 X 2-2, 5

mm, glandular-villous; verticillasters 4-6-flowered,

2-3,5 cm apart. Calyx 9 mm long at anthesis, glan-

dular-villous; upper tooth broadly ovate, rounded,

2,5 mm long, decurrent; lower 4 teeth deltoid-

subulate, the lowermost pair the longest, up to 2,5 mm

long. Corolla 14 mm long, minutely puberulous;

tube 10 mm long, widening from the middle to 4—5 mm

wide at the mouth; upper lip formed by the oblique

mouth of the corolla with a minute lobe; lower lip

concave, 4 mm long, usually deflexed. Stamens

exserted by 10 mm, far exceeding the lower lip;

upper pair attached below the middle of the tube,

filaments minutely puberulous below; upper pair

attached at the throat, filaments united to near the

apex.

A mountain grassland species known from only

one gathering at 3 000 m in the Natal Drakensberg.

Natal. — 2929 (Underberg): near Giants Castle (-BC),

A. Bolus sub Guthrie 4894 (BOL).

The habit and corolla shape suggest a relationship

to H. transvaalensis (Schltr.) Ashby, but the leaves

of the latter are acute and markedly toothed while

the apex of the inflorescence is adorned by a coma

of colourful bracts.

N. E. Brown, l.c., quotes this species as affording

an example of the lower pair of filaments being free

or united on the same plant. There are two capsules

mounted on the type specimen, one marked “A,

flower with filaments free” and the other “B, flower

with lower pair of stamens united nearly to the apex”.

It must be concluded that Flower A must have been

abnormal or badly squashed in pressing. There are

several good flowers on the sheet and in at least six

of them the lower pair of filaments are obviously

united to near the apex without the need for dissecting

them. More material of the species is desired for

further study.

19. Hemizygia ramosa Codd, sp. nov. a H. trans-

vaalensis (Schltr.)Ashby habitu ramosissimo, foliis

obscure dentatis, verticillastris 2-floribus differt.

Frutex ramosissima, 1-1 ,2 m altus; ramuli breviter

tomentosi. Folia breviter petiolata; petiolus 1-3 mm

longus, tomentosus; lamina obovata vel oblanceolata,

15-25 mm longa, 6-1 1 mm lata, supra parce hispidula,

subtus hispidula, glanduloso-punctata, apice rotun-

dato, basi obtusa vel cuneata, margine praecipue

supre medium obscure crenato-dentata. Inflorescentia

plerumque basin versus parce ramosa, 10-15 cm

longa; rhachis parce strigilosis; bracteae ovatae vel

late ellipticae, 14-16 mm longae, 6-8 mm latae,

lilacinae, subglabrae vel parce pubescentes; verti-

cillastri 2-flori; pedicelli 2-3 mm longi. Calyx per

anthesin 8 mm longus, parce glanduloso-hispidulus ;

tubus 6 mm longus; lobus posticus late ovatus,

rotundatus, 2 mm longus, margine decurrenti; dentes

laterales deltoideo-subulati, 1 mm longi; antici

subulati, 2 mm longi. Corolla malvina, 25-28 mm

longa, puberula; tubus 20-22 mm longus apicem

versus sensim ampliatus, ore 4-5 mm lato; labium

posticum 2 mm longum; anticum concavum, horizon-

tale vel recurvum, 5 mm longum. Stamina 9-1 1 mm

exserta; postica circa medium tubi corollae inserta,

filamentis libris, basin versus pubescentibus ; antica

fauce corollae inserta, filamentis ad apicem connatis.

Stylus 15 mm exsertus, apice breviter bilobato.

Type: Natal, 2732 (Ubombo), near Mkuze (-CA),

Moll 3158 (PRE, holo.).

Shrub, much branched, 1-1,2 m tall; branchlets

shortly tomentose. Leaves shortly petiolate; petiole

1-3 mm long, tomentose; blade obovate to oblan-

ceolate, 15-25 mm long, 6-11 mm broad, sparingly

hispid above, hispid and gland-dotted below, apex

rounded, base obtuse to cuneate, margin obscurely

crenate-dentate mainly above the middle. Inflorescence

usually sparingly branched towards the base, 10-15

cm long; rhachis sparingly strigilose; bracts ovate to

broadly elliptical, 14-16 mm long, 6-8 mm broad,

mauve-pink subglabrous to sparingly pubescent;

verticillasters 2-flowered; pedicels 2-3 mm long.

Calyx when flowering 8 mm long, sparingly glandular-

hispidulous; tube 6 mm long; posticous lobe broadly

ovate, rounded, 2 mm long, margin decurrent;

lateral teeth deltoid-subulate, 1 mm long; anticous

teeth subulate, 2 mm long. Corolla mauve, 25-28 mm

long, puberulous; tube 20-22 mm long, widening

gradually towards the apex, mouth 4-5 mm wide;

posticous lip 2 mm long; anticous lip concave,

horizontal or recurved, 5 mm long. Stamens exserted

by 9-11 mm; posticous stamens attached about the

middle of the corolla tube, filaments free, pubescent

towards the base; anticous stamens inserted in the

throat of the corolla, filaments united to the apex.

Style exserted by 15 mm, apex shortly bilobed.

Fig. 7.

Found in shallow soil among rocks in open wood-

land at the southern end of the Lebombo Range

near Mkuze.

Natal. — 2732 (Ubombo): Lebombo Mts., near Mkuze

(-CA), Moll 3158; Mkuze Poort (-CA), Ward 4074.

In the herbarium, H. ramosa is reminiscent of

H. transvaalensis (Schltr.) Ashby but the two are not

likely to be confused in the field because of differences

in habit and ecology. H. ramosa is a much-branched

bushy shrub growing on rocky, wooded hillsides and

cliffs, while H. transvaalensis is adapted to grassland

which is periodically burnt, with the result that it

develops a woody rootstock from which several

branches arise annually. There are also differences

in shape, pubescence and nervation of leaves, with

H. ramosa having leaves more obovate and less dentate

than those of H. transvaalensis. An important

difference, in addition, are the flowers produced

singly in the axils of the bracts in H. ramosa while,

in H. transvaalensis each bract subtends usually

three flowers.

In the field H. ramosa would probably remind one

of Syncolostemon latidens (N.E. Br.) Codd, thus

emphasizing the close affinity of the two genera.

However, the calyx of H. ramosa is clearly of the

Hemizygia type, having a broad upper lobe, decurrent

on the tube, with the remaining four teeth ending in

subulate, almost spine-like points.

L. E. CODD

15

Fig. 7. — Hemizygia ramosa Codd ( Moll 3158, PRE, holotype).

20. Hemizygia transvaalensis ( Schltr .) Ashby in J.

Bot. Lond. 73: 349 (1935); Letty, Wild Flow. Transv.

285, 1. 141 , 4 (1962). Type: Transvaal, Barberton,

Galpin 468 (PRE; SAM).

Orthosiphon transvaalensis Schltr. in J. Bot. Lond. 35: 281

(1897); N.E. Br. in FI. Cap. 5,1 : 244 (1910). O. wilmsii Guerke

in Bot. Jahrb. 26: 79 (1898). Syntypes: Transvaal, Lydenburg,

Wilms 1107 (BM; K; PRE); 1108. O. muddii N.E. Br. l.c. 245

(1910). Syntypes: Transvaal, Drakensberg, Mudd s.n. (K;

PRE, fragment); Spitzkop, Burtt Davy 1570 (K).

Soft shrublet 30-100 cm tall; stems arising annually

from a perennial woody rootstock, becoming woody

at the base, sparingly to freely branched and broom-

like (in the latter case with many small leaves),

sparingly to densely hispid. Leaves sessile or shortly

petiolate; blade stiff, on main stems ovate to broadly

ovate, 1 5—40 mm long, 8-22 mm broad, on branched

form ovate-elliptic to ovate, 12-20 mm long, 4-8 mm

broad, pale greenish-brown and veins impressed

above, slightly paler and reticulate below, sparingly

to densely pubescent and glandular on both surfaces,

especially on the nerves below; apex acute to obtuse,

base obtuse to rounded; margin fairly coarsely

serrate-dentate for the upper two thirds, rarely teeth

obscure. Inflorescence 7-20 cm long, fairly lax,

simple or with 1 or 2 pairs of branches near the base;

rhachis glandular-hispid; bracts ovate to lanceolate,

the terminal ones, persistent, pairs spaced 10-15 mm

apart, pinkish-purple, 12-24x4-10 mm, subglabrous;

verticillasters usually 6-flowered, 1-2 cm apart.

Calyx 10-12 mm long at anthesis, densely glandular-

hispidulous, purple; upper lip ovate, rounded,

2,5 mm long, decurrent; lower 4 teeth deltoid-subulate

to bristle-like, the lowest pair the longest, up to 3 mm

long. Corolla whitish to mauve-purple or lilac-pink,

1 8—22 mm long, glabrous; tube 14-17 mm long

widening to 5 mm wide at the mouth; upper lip a

small appendage; lower lip concave, 4-6 mm long,

often deflexed. Stamens exserted by 7-10 mm,

exceeding the lower lip of the corolla; upper pair

attached about the middle of the tube, filaments

pubescent in the lower two thirds; lower pair attached

at the throat, filaments united nearly or to the apex.

Stigma bifid.

Locally common at medium altitudes of 1 000 to

1 700 m in the eastern Transvaal mountains from

the Mariepskop and Lydenburg areas southwards to

Barberton, on grassy slopes and flats, usually among

rocks.

Transvaal. — 2430 (Pilgrim’s Rest): near Mariepskop (-DB),

Fitzsimons & Van Dam sub TR V 26256; Van der Schijff 5888;

Bourkes Luck Mine (-DB), Galpin 14313; Ohrigstad Nature

Reserve (-DC), Jacobsen 1468; Lisbon Falls (-DD), Jordaan

99; Pilgrim’s Rest (-DD), Rogers 14373; 18251; Galpin 14553;

1 8 km E. of Graskop, Codd & de Winter 3123. 2530 (Lydenburg) :

16 km N. of Lydenburg (-AB), Strey 4113; near Lydenburg

(-AB), Wilms 1107; Young A452; Codd 539; farm Rietfontein

1240 (-AB?), Burtt Davy 7256; farm Zwagershoek (-AB),

Obermeyer 328; Obermeyer & Verdoorn 32; farm Kleinfontein

(-AC), Burger 13; 26 km S.E. of Lydenburg (-BA), Morris 21;

10 km W. of Sabie (-BA), Balsinhas & Kersberg 2139; 11 km E.

of Sabie (-BB), Brent 114; Crocodile River (-BC?), Schlechter

3916; Wonderkloof Nature Reserve (-BC), Kluge 24; Elan-

Puttick 47; Rosehaugh (-BD), Mogg 17571; near Nelspruit

(-BD), Rogers sub TRV 4741; Buitendag 557; 18 km S.E. of

Sewefontein (-CD), Codd 8112; Godwan River (DA), Prosser

1264; Coetzeestroom Forestry Station (-DA), Hardy 11;

Kaapschehoop (-DB), Thode A 1634. 2531 (Komatipoort):

Pretorius Kop (-AB), Lang sub TRV 31557 ; White River (-AC);

Thorp sub NH 29672; near Barberton (-CC), Galpin 468;

Bolus 7604; Thorncroft sub TRV 3123; Williams sub TRV 7655;

Rogers 21442; Mauve 4806.

H. transvaalensis is related to the next two species,

H. foliosa S. Moore and H. thorncroftii (N.E. Br.)

Ashby, but is a more robust species, up to 1 m tall,

with usually broader bracts and longer flowers

(corolla tube 14-17 mm long). Depauperate specimens

may be only 20 cm tall with narrow bracts and

these may be confused with H. thorncroftii which,

however, usually has narrowly elliptical leaves while

the corolla tube is 8-10 mm long. H. foliosa tends to be

decumbent, usually with large, elliptical leaves

rounded at the apex, the corolla is smaller (tube

about 10-12 mm long), and the verticillasters are

2-flowered.

Occasional specimens of H. transvaalensis branch

freely and produce numerous small leaves, giving

specimens a broom-like appearance. Such a specimen

was described as Orthosiphon wilmsii Guerke.

However, there are many intermediates linking it

with typical H. transvaalensis in which branching is

sparing and leaves fewer and larger. There is also a

good deal of variation in leaf shape, toothing of the

margin and pubescence. It grows under warmer and

drier “middleveld” conditions than the majority of

eastern Transvaal species and the showy bracts and

flowers have led to it being cultivated with some

success in Pretoria.

21. Hemizygia foliosa S. Moore in J. Bot. Lond.

43: 172 (1905); Ashby in J. Bot. Lond. 73: 348 (1935);

Compton, FI. Swaz. 67 (1966). Type: Swaziland,

Mbabane, Burtt Davy (BM, holo.; K; PRE).

Orthosiphon foliosus (S. Moore) N.E. Br. in FI. Cap. 5,1:

243 (1910). O. humilis N.E. Br., l.c. 259 (1910). Type: Transvaal,

Waterval Onder, Rogers 4375 (K, holo.; PRE).

Hemizygia humilis (N.E. Br.) Ashby, l.c. 348 (1935). Type as

above.

16

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

Herbaceous stems from a perennial woody root-

stock; stems 1-several, erect to decumbent 20-35 cm

long, softly woody at the base, thinly to densely

villous. Leaves sessile or very shortly petiolate; blade

ovate to ovate-elliptic or elliptic, varying but usually

large when mature, 2,5-7 cm long, 1,5-3, 5 cm

broad, concolorous, puberulous and somewhat shiny

to appressed pilose, gland-dotted on both surfaces;

apex obtuse to rounded, base obtuse to truncate;

margin entire to somewhat distantly dentate.

Inflorescence paniculate with 2 or 3 pairs of branches

near the base or occasionally simple, lax, 10-20 cm

long; rhachis glandular-hispid to villous; bracts

deciduous below, usually persisting as a purplish coma

at the apex, small to fairly large, ovate-lanceolate,

8-18x3-8 mm; verticillasters 2-flowered, 1-2 cm

apart. Calyx 7-8 mm long at anthesis, glandular-

hispid; upper tooth broadly ovate, 2 mm long,

decurrent; lower 4 teeth shortly deltoid-subulate the

lower pair the longer, up to 2 mm long. Corolla

whitish to mauve, 12-14 mm long, puberulous on the

lips; tube 9-10 mm long, widening to 4 mm at the

mouth; upper lip a small appendage; lower lip

concave, 3-4 mm long, often deflexed. Stamens

exserted by 10-12 mm, well exceeding the lower lip;

upper pair attached about the middle of the tube,

filaments pubescent below; lower pair attached at

the throat, filaments united to the apex. Stigma

bifid.

Found in dense mountain grassland, often among

rocks, at altitudes of 1 300 to 1 700 m in the south-

eastern Transvaal and western Swaziland.

Transvaal. — 2530 (Lydenburg): Waterval Onder (-CB),

Rogers 4735; Nelshoogte (-DD), Prosser 1466. 2630 (Carolina):

Arnhemburg (not located), Roberts sub TRV 15853. 2731

(Louwsburg): 30 km W. of Pongola Settlement (-AD ),Codd2101

Swaziland. — 2631 (Mbabane): near Mbabane(-AC), Burtt

Davy 2833; Bolus 12250; Compton 25241; 25271; Usutu

Forests Concession (-AC), Codd 9507; near Mankaiana (-CA),

Codd 4726; Sidey 1928; Hlatikulu (-CD), Galpin 10207.

May be distinguished from H. transvaalensis and

H. thorncroftii by the 2-flowered verticillasters and

the large, ovate to ovate-elliptic, entire or distantly

toothed leaves. In some specimens the leaves are

only 2 cm long, but these are probably immature.

There is also a good deal of variation in pubescence,

from puberulous to pillose leaves. The terminal

bracts also vary in size, in some specimens being less

than 1 cm long, but are always purplish in colour.

The type of H. humilis is such a specimen and the

leaves are somewhat smaller than usual but the modern

range of material links it with H. foliosa.

22. Hemizygia thorncroftii (N.E. Br .) Ashby in

J. Bot. Lond. 73: 349 (1935); Compton, FI. Swaz.

67 (1966). Lectotype: Transvaal, Barberton, Thorn-

croft 3123 (K, lecto.).*

Orthosiphon thorncroftii N.E. Br. in FI. Cap. 5,1 : 246 (1910).

Herb 15-30 cm; stems several arising annually

from a perennial woody rootstock, subglabrous to

glandular-hispid. Leaves subsessile; blade elliptic to

linear-elliptic, 15-40 mm long, 4-10 mm broad,

concolorous, sparingly pubescent on both surfaces;

apex obtuse to acute, base cuneate to attenuate;

margin with a few small distant teeth near the apex.

Inflorescence simple or occasionally with a pair of

branches near the base, 7-20 cm long, lax; rhachis

glandular-hispid; bracts deciduous below, the upper

ones persisting as a purple coma, lanceolate to

linear-lanceolate, 15-30x2-5 mm; verticillasters

usually 6-flowered, 1-2,5 cm apart. Calyx 8-9 mm

* Ashby points out that, in the Transvaal Museum, No. 3125

is H. thorncroftii while No. 3123 is H. transvaalensis.

long at anthesis, glandular-hispid; upper tooth ovate,

3 mm long, decurrent; lower 4 teeth deltoid-subulate,

the lowest pair the longest, up to 3 mm long. Corolla

mauve, 14—16 mm long, puberulous on the lips;

tube 10-12 mm long, widening to 4 mm at the mouth;

upper lip a small appendage; lower lip concave, 4 mm

long, often deflexed. Stamens exserted by 10 mm, well

exceeding the lower lip; upper pair attached about

the middle of the tube or just below, filaments

pubescent in the lower part; lower pair attached at

the throat, filaments united for their entire length.

Stigma minutely bifid.

Found on grassy slopes at altitudes of 1 000 to

1 800 m in the Barberton District of Transvaal and

in western Swaziland.

Transvaal. — 2531 (Komatipoort): near Barberton (-CC),

Thorncroft sub TRV 3125; Galpin 465; Codd 9791; Lomati

Valley (-CC), Thorncroft 2015.

Swaziland. — Without locality, Stewart sub TRV 12737.

2631 (Mbabane): near Mbabane, Compton 25222; 26050;

26939; 28090; 29078; 30878; near Mankaiana, Compton

29165; Hardy 1623.

Although resembling depauperate specimens of

H. transvaalensis, H. thorncroftii can usually be

recognized by the long-elliptical leaves and the long,

narrow apical bracts, while the shorter corolla tube

(10-12 mm as against 14-17 mm in H. transvaalensis)

will be diagnostic.

Individual flowers resemble those of H. modesta but

the latter species has 2-flowered verticillasters and

lacks the colourful apical bracts of H. thorncroftii.

23. Hemizygia persimilis (N.E. Br.) Ashby in J.

Bot. Lond. 73: 349 (1935). Lectotype: Transvaal,

Barberton, Thorncroft sub TRV 3132 (K, lecto.;

PRE; SAM).

Orthosiphon persimilis N.E. Br. in FI. Cap. 5,1 : 246 (1910).

O. rogersii N.E. Br., l.c. 247 (1910). Syntypes: Nelspruit,

Rogers sub TRV 4740 (K; PRE; SAM); Devil’s Kantoor,

Kaapschehoop, Bolus 9742.

Bushy herb 15-30 cm tall from a perennial root-

stock; stems simple or branched, glandular-hispid.

Leaves subsessile; blade lanceolate-elliptic, 15-20 mm

long, 7-9 mm broad, more or less concolorous,

sparingly appressed-pubescent, the surface some-

what wrinkled and dotted with brownish gland-dots;

apex obtuse, base cuneate ; margin entire. Inflorescence

simple, 3-12 cm long, fairly dense; rhachis densely

glandular-villous; bracts both the lower and upper

persistent, the upper ovate-lanceolate to broadly

ovate, 12-15x7-10 mm, whitish to rose-purple,

thinly pubescent and dotted with reddish-brown

gland-dots; verticillasters 2-6-flowered, 3-10 mm

apart. Calyx 8 mm long, glandular-villous and freely

gland-dotted; upper tooth ovate, rounded, decurrent;

lower 4 teeth deltoid-subulate, the lower pair the

longer, up to 2 mm long. Corolla white, drying

yellow-brown, 11-12 mm long, puberulous both on

the outside and within the upper lip; tube 8 mm long,

not expanding towards the throat, lips spreading;

upper lip formed by the abruptly spreading throat,

erect, 3,5 mm long; lower lip concave, 3-4 mm long.

Stamens shortly exserted by 1-2 mm, the upper pair

sometimes scarcely exserted; upper pair attached

about 1 mm from the base of the corolla tube, fila-

ments pubescent for their entire length; lower pair

attached at the throat, joined at their point of insertion

or united only at the base for ca. 0,5 mm, filaments

sparingly pubescent. Stigma clavate.

Known only from the Nelspruit-Barberton area

of the eastern Transvaal where it grows in grassy

places among rocks in lowveld woodland at altitudes

of about 1 000 m.

L. E. CODD

17

Transvaal.— 2530 (Lydenburg); near Nelspruit (-BD),

Rogers sub TRV 4740; Holt 300; Mauve 4942; Prosser 1489

(J); Buitendag 166; 582 (NBG). 2531 (Komatipoort): near

Barberton, Thorncroft sub TRV 3132.

H. persimilis shows a combination of unusual

features in the genus, some of which are also found

in H. pretoriae, a species of similar stature and

ecology. For example, the narrow corolla tube

expands abruptly at the mouth forming what appears

to be a longish upper lip nearly equal in length to the

lower lip, and the stamens are very shortly exserted

(the upper pair is included in H. pretoriae and

exserted by 1-2 mm in H. persimilis). In addition,

H. persimilis differs in the upper pair of stamens being

attached very near the base of the corolla (about one

third up the tube in H. pretoriae ); the lower pair of

filaments may be free or united for only about 0,5 mm

at the base (united for most of their length in H.

pretoriae)', the stigma is clavate (slender and shortly

bifid in H. pretoriae)', and the bracts are large and

showy (small and inconspicuous in H. pretoriae).

There are thus as many differences as there are simila-

rities between the two species, both of which are rather

aberrant members of the genus.

The fact that the filaments of the lower pair of

stamens are united only at the base may be put

forward as an example which breaks down the

distinction between Hemizygia and Orthosiphon.

However, the structure is different. In Orthosiphon

the filaments (free to the point of attachment) continue

down as distinct and separate ridges on the corolla

tube; in H. persimilis the filaments are united at their

point of attachment and continue as a single confluent

ridge down the corolla-tube.

24. Hemizygia petiolata Ashby in J. Bot. Lond.

73: 355 (1935). Type: Transvaal, Soutpansberg,

Tshakoma, Obermeyer sub TRV 31571 (PRE, holo.).

Soft shrub up to 1 m tall, branching usually from

the base; stems few to many, ascending, glandular-

pilose. Leaves petiolate; petiole 6-14 mm long; blade

ovate to ovate-lanceolate, 2-5,5 cm long, 0,6-3 cm

broad, dark brown and shortly glandular pubescent

above, paler and canescent below, nerves distinct;

apex acute to obtuse, base cuneate to obtuse; margin

regularly serrate-dentate in the upper two thirds.

Inflorescence paniculate or occasionally simple, lax,

10-30 cm long; rhachis glandular-tomentulose; bracts

sometimes persisting as a purple coma, ovate, usually

rather small, 5-10x3-5 mm, more often the apex of

the inflorescence broken off, lower bracts early

caducous ; verticillasters (4-) 6-flowered, 1 , 5^4 cm

apart. Calyx 5-6 mm long at anthesis, enlarging in

fruit, glandular-tomentose; upper tooth ovate to

subrotund, 1,5-2 mm long, decurrent; lower 4 teeth

deltoid-subulate to bristle-like, up to 2 mm long.

Corolla pale mauve to lilac, 17-20 mm long, finely

pubescent; tube 13-16 mm long, narrowly cylindrical

for about 8-10 mm then expanding to 3-4 mm at the

mouth; upper lip a small appendage; lower lip

concave, 4 mm long. Stamens exserted by 7-10 mm,

exceeding the lower lip; upper pair attached about

3 mm from the base of the tube, filaments puberulous

below; lower pair attached at the throat, filaments

united for more than half their length. Stigma swollen,

emarginate.

Restricted to north-eastern Transvaal from the

Soutpansberg to near Duiwelskloof on rocky, wooded

hillsides and at forest margins, at altitudes of 1 000 to

1 600 m.

Transvaal. — 2229 (Waterpoort): Hanglip (-DD), Meeuse

10171; Punch Bowl Hotel (-DD), Codd 8331. 2230 (Messina):

Entabeni Forest Station (-CC), Bruce <& Kies 78; Codd 8394;

Sibasa (-CD), Junod sub TRV 25470; Munro s.n. 2329 (Peters-

burg): near Louis Trichardt (-BB), Breyer sub TRV 22727;

Ihlenfeldt 2216. 2330 (Tzaneen): Tshakoma (-AB), Obermeyes

sub TRV 31571; Westfalia Estate (-CA), Scheepers 387; Bor

1355; Woodbush (-CC), Bruce dk Kies 78; Codd 9423. 2430

(Pilgrim’s Rest): The Downs (-AA), Junod 4358.

A strongly aromatic plant allied to the next species,

H. canescens (Guerke) Ashby, but has more ovate

leaves, longer petioles and longer corolla tube.

25. Hemizygia canescens ( Guerke ) Ashby in J.

Bot. Lond. 73: 354 (1935); Compton, FI. Swaz. 67

(1966); Ross, FI. Natal 306 (1972). Lectotype:

Transvaal, Wonderboompoort, Rehmann 4507 (Z,

lecto.; K).

Orthosiphon canescens Guerke in Bull. Herb. Boiss. 6:

557 (1898); N.E. Br. in FI. Cap. 5,1 : 259 (1910). O. affinis N.E.

Br., l.c. 257 (1910). Syntypes: Transvaal, Woodbush Mts.,

Schlechter 4737 (K; PRE); near Potgietersrus, Bolus 11146

(BOL).

Herb, probably perennial, woody and branched

below, 30-60 cm tall; stems often branched, shortly

greyish-tomentose, often crisped or, occasionally,

sparse but not villous. Leaves subsessile or shortly

petiolate; petiole up to 5 mm long; blade linear or

linear-lanceolate to lanceolate or, rarely ovate-

lanceolate, 2, 5-5, 5 cm long, 3-15 mm broad, con-

colorous, densely canescent on both surfaces to

sparingly short crisped tomentulose and somewhat

rugose, nerves prominent below; apex acute, base

cuneate to attenuate; margin finely to fairly coarsely

serrate-dentate in the upper half to two thirds.

Inflorescence simple to freely branched, 7-25 cm long,

lax; rhachis crisped-tomentulose to finely glandular-

hispidulous; bracts early deciduous, small, ovate,

ca. 2x1 mm; verticillasters 4-6-flowered, 1-3 cm

apart. Calyx 5 mm long at anthesis, enlarging in

fruit, glandular-tomentulose to hispidulous; upper

lip ovate to subrotund, 2 mm long, decurrent; lower

4 teeth deltoid-subulate, becoming bristle-like, up to

2 mm long. Corolla white or pale mauve or purplish,

14-17 mm long, finely pubescent; tube 10-13 mm long,

widening abruptly about 2,5 mm from the throat to

3-4 mm wide at the throat; upper lip a small appen-

dage; lower lip concave, 3-4 mm long. Stamens

exserted by 10 mm, well exceeding the lower lip; upper

pair attached about 4 mm from the base of the tube,

filaments puberulous below; lower pair attached at

the throat, filaments united for most of their length.

Stigma somewhat clavate.

Distributed in a broad band from the Mafeking

District of the Cape Province, across south-western

and central Transvaal to eastern Transvaal, avoiding

the high mountains, extending to Swaziland and

northern Zululand; among rocks in open arid to moist

woodland and marginal grassland at altitudes of

300 to 1 700 m.

Transvaal. — 2329 (Pietersburg) : Houtboschberg (-DD),

Schlechter 4737; 4797. 2330 (Tzaneen): Hans Merensky Nature

Reserve (-CC), Gilliland 764; 16 km S. of Tzaneen (-CC),

Bruce & Kies 77. 2427 (Thabazimbi) : between Hermanusdooms

and Elmeston (-BA), Meeuse & Strey 10427. 2428 (Nylstroom):

Geelhoutkop (-AD), Breyer sub TRV 17781; Mosdene, near

Naboomspruit (-DA), Galpin M 286. 2429 (Zebediela): Pyramid

Estate near Potgietersrus (-AA), Galpin 9067; 9 km N.W. of

Marble Hall (-CD), Codd 10368. 2526 (Zeerust): near Zeerust

(-CA), Jenkins sub TRV 11692. 2527 (Rustenburg): near

Rustenburg (-CA), Collins 30; Jacobsen 836; Silikaatsnek

(-DB), Acocks 12430. 2528 (Pretoria): Hooms Nek (-CA),

Verdoorn s.n.; near Pretoria (-CA), Leendertz sub TRV 8555;

Mogg 16373; 16475; Donkerhoek (-CB), Repton 1311. 2529

(Witbank): Loskopdam (-AD), Codd 9840; Theron 1671;

Doornkop 273 (-CB), Du Plessis 348; Botsabelo (-CB), Schlech-

ter 4070. 2530 (Lydenburg): 8 km E. of Sabie (-BB), Eliovson

sub J38580 (J); Witklip (-BD), Kluge 434; near Nelspruit

(-BD), Breyer sub TR V 17909; Buitendag 438 (NBG) ; Kaapsche

Hoop (-DB), Rogers 20823. 2531 (Komatipoort): Plaston

(-AC), Holt 198. 2726 (Potchefstroom): near Potchefstroom

18

THE SOUm AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

(-CA), Potts s.n.; Lonw 699. 2628 (Johannesburg): near Heidel-

berg (-AD), Leendertz 1027 ; Repton 824; Story 1606; Suiker-

bosrand (-CA), Bredenkamp 762.

Swaziland. — 2631 (Komatipoort): Piggs Peak (-CC),

Compton 27626; 27657. 2631 (Mbabane): Stegi, Blue Jay

Ranch (-BD ?), Compton 31458.

Natal. — 2732 (Ubombo): Pongolo Poort (-AC), Ward

4083.

Cape.— 2525 (Mafeking): near Mosita (-DC?), Brueckner

s.n.; 538.

A good deal of variation in leaf shape is included

in H. canescens from linear (3-4 mm wide), in the

dry western extremity of its range in the Mafeking

District, to lanceolate and ovate-lanceolate (up to

15 mm wide) in more mesophytic areas. The species is

diagnosed on the basis of the short canescent, often

crisped tomentum of stems and leaves though,

towards the north of the range, in the Waterberg,

Potgietersrus and Tzaneen Districts, the pubescence

is more scanty and somewhat rougher. This form

was separated as Orthosiphon affinis N.E. Br., but

Ashby reduced it to synonymy under H. canescens.

Although the extremes can be separated with close

scrutiny under magnification, there are numerous

intermediates linking them.

Supperficially H. canescens closely resembles H.

petrensis but the latter may be recognized by the

presence of long villous hairs on the stems, although the

pubescence of the leaves is often similar. The latter is

a more western species, entering the northern and

eastern Transvaal lowveld, and is also very variable

(see p. 19). H. canescens appears to be a fairly clear-

cut entity with a distribution distinct from H. petrensis

and thus it seems justified to uphold both as species.

However, two specimens from the Waterberg in S.W.

Africa, Boss sub TRV 35003 and De Winter 2799,

have pubescence resembling H. canescens and this

areas should be investigated further.

26. Hemizygia linearis ( Benth .) Briq. in Bull.

Herb. Boiss. ser. 2,3: 997 (1903); Ashby in J. Bot.

Lond. 73: 354 (1935). Type: Matabeleland, Oates s.n.

(K, holo.).

Orthosiphon linearis Benth. in Hook. Ic. PI. t. 1274 (1878);

Rolfe in Oates, Matabeleland ed. 2: 407 (1889); Bak. in FI.

Trop. Afr. 5: 374(1900).

Herb, probably perennial, 30-50 cm tall, somewhat

woody and branching below; stems subglabrous to

sparingly villous, quadrangular and ribbed along the

angles. Leaves sessile or subsessile; blade linear, 2-3

cm long, 2^4 (-5) mm broad, puberulous to sparingly

hispid, often folded along the midrib or with margins

inrolled; apex acute, base attenuate; margin finely

and distantly toothed. Inflorescence 1 2—20 cm long,

simple or branched near the base, lax; rhachis

sparingly hispidulous; bracts early caducous, very

small, ovate, 2x1 mm, verticillasters 4-6-flowered,

1 , 5-4 cm apart. Calyx 5 mm long at anthesis,

hispidulous; upper tooth broadly ovate, rounded,

2 mm long, purple, decurrent; lower 4 teeth deltoid-

subulate, up to 1,5 mm long, becoming bristle-like.

Corolla mauve, often with violet stripes, 12-13 mm

long, finely pubescent; tube 9-10 mm long widening

abruptly about 3 mm from the throat to 3 mm wide

at the throat; upper lip 1,5 mm long; lower lip

concave, 3 mm long. Stamens exserted by 6-8 mm,

exceeding the lower lip; upper pair attached about

3 mm from the base of the tube, filaments puberulous

below; lower pair attached at the throat, filaments

united nearly to the apex. Stigma somewhat clavate.

Found in open places in dry woodland in South

West Africa and northern Cape Province; also in

Rhodesia and Angola.

S.W. A.— 1821 (Andara): Andara Mission Station (-AB),

De Winter <£- Marais 4789. 2217 (Windhoek): Auas Mts. (-CA)’

Strey 2571; farm Lichtenstein (-CC), Merxmuller dk Giess

1247; farm Rietfontein (-CD), Strey 2564.

Cape. — 2723 (Kuruman): Takoon (-BB), Burtt Davy 13961 .

Diagnostic features are the linear, subglabrous

leaves and the subglabrous to sparingly villous stems

which have a somewhat varnished appearance.

H. petrensis is closely related to it and, as may be

expected, some specimens are difficult to allocate with

certainty, but H. petrensis usually has a strong

development of villous hairs on the stems and, to a

lesser extent, on the leaves (see also below). Some

specimens of H. canescens have linear leaves but the

dense, short pubescence on stems and leaves can be

used to exclude such specimens from H. linearis.

27. Hemizygia petrensis (Hiern) Ashby in J. Bot.

Lond. 73: 353 (1935); Launert & Schreiber in Prodr.

FI. S.W. Afr. 123: 13 (1969). Type: Angola, Wel-

witsch 5494 (BM, holo.).

Orthosiphon petrensis Hiern, Cat. Afr. PI. Welw. 1 : 859

(1900); Bak. in FI. Trop. Afr. 5: 524 (1900). O. dinteri Briq.

in Bull. Herb. Boiss. ser. 2,3: 995 (1903). Type: 10 km E. of

Orumbe, Dinter 1320. O. varians N.E. Br. in FI. Cap. 5,1 : 256

(1910); Ashby, l.c. 357 (1935). Type: Transvaal, Komatipoort,

Schlechter 11746 (BOL!, holo.). O. holubii N.E. Br., l.c. 258

(1910). Type: Molopo River, Holub s.n. (K, holo.). O. engleri

Perkins in Bot. Jahrb. 54: 344 (1917). Type: S.W. Africa,

Okahandja, Engler 6475. O. mossianus Good in J. Bot. Lond.

63: 175 (1925). Type: Transvaal, Messina, Moss & Rogers

193 (BM, holo.; PRE).

Hemizygia mossiana (Good) Ashby, l.c. 356 (1935).

Strongly aromatic herb, annual or perennial,

20-60 cm tall, branching near the base and woody

below, with a woody taproot; stems villous to densely

villous with long, spreading greyish-white hairs,

quadrangular and often strongly ribbed along the

angles. Leaves subsessile or shortly petiolate; blade

variable from linear-lanceolate to oblong-lanceolate

or ovate-lanceolate 2-5 cm long, 5-15 mm broad,

sparingly to densely pilose or canescent, often with

long and short hairs intermingled; apex acute, base

cuneate to attenuate; margin obscurely to distinctly

and somewhat distantly toothed. Inflorescence 8-20

cm long, lax, simple or with a pair of branches near the

base; rhachis glandular-villous; bracts caducous,

small, ovate to broadly ovate, 3 mm x 2 mm,

pubescent; verticillasters 4-6-flowered, 1-3 cm apart.

Calyx 4-5 mm long at anthesis, enlarging in fruit,

glandular-hispid to villous; upper lip broadly ovate

or subrotund, purple, 2 mm long, decurrent; lower

4 teeth deltoid-subulate, 1 , 5 mm long. Corolla

pinkish to lilac or voilet, finely pubescent, 13-15 mm

long; tube 9-12 mm long, widening abruptly about

3 mm from the apex to 2,5-3 mm wide at the throat;

upper lip a small appendage; lower lip concave,

3-4 mm long. Stamens exserted by 8 mm, exceeding

the lower lip; upper pair attached 2-3 mm from the

base of the tube, filaments puberulous below; lower

pair attached at the throat, filaments united for more

than two thirds their length. Stigma somewhat

clavate.

Recorded from northern South West Africa and

northern and eastern Transvaal, among rocks, in

open places and watercourses in semi-arid woodland

at altitudes of 200-700 m in the Transvaal and up

to 2 000 m in the Windhoek area of South West

Africa. Also in Angola and Rhodesia.

S.W. A. —1917 (Tsumeb): Otavifontein (-CB), Dinter 5305;

Guchab (-DB), SchoenJ elder 945. 2017 (Waterberg): Waterberg

Plateau (-AC), Boss sub TRV 35003; De Winter 2799. 2217

(Windhoek): farm Regenstein (-CA), Giess 11675.

L. E. CODD

19

Transvaal. — 2229 (Waterpoort): farm Greefswald (AB),

Codd d Dyer 3832; Bruce 54; Pienaar 306; 340; Zoutpan (-CD),

Schweickerdt d Verdoorn 613. 2230 (Messina): near Messina

(-AC), Moss d Rogers 193; Rogers 20811 ; 22494; Wild 7631.

2231 (Pafuri): Makuleke’s Location (-AC), Obermeyer 656;

Baiandbai (-CC?), Lang sub TRV 32214; 32228. 2329 (Peters-

burg): near Sand River (-BA), Meeuse 10211. 2330 (Tzaneen):

Hans Merensky Nature Reserve (-CC), Zambatis H2. 2331

(Phalaborwa): Kruger National Park, Letaba Bridge (-DC),

Van der Schijff 545; 547; The Gorge (-DD), Van der Schijff

2331. 2430 (Pilgrim’s Rest): Harmony Block, farm Calais

(-AB?), Breyer sub TRV 25216; 16 km N. of Abel Erasmus

Pass (-BC), Mauve 4331. 2431 (Acornhoek): Kruger National

Park, between Skukuza and Tshokwane (-DD), Codd d De

Winter 5073; Schlieben 9392. 2531 (Komatipoort): near Komati-

poort (-BD), Schlechter 11746 (BOL); Hilliard d Burtt 3632;

junction of Sigaas and Crocodile Rivers (-CB), Van der Schijff

3951.

The three species H. canescens, H. linearis and H.

petrensis form a closely related group with almost

identical floral characters and small, inconspicuous

bracts. H. canescens may be dinstinguished on the

basis of the dense, short and often crisped pubescence

on stems and leaves and is distributed mainly on the

high plateau formed by the northern Cape, south-

western and central Transvaal, extending to eastern

Transvaal, Swaziland and Natal. In H. linearis, which

is the oldest name, the leaves are linear to filiform

(occasionally some leaves up to 5 mm broad) and

leaves and stems are subglabrous or with scattered

long hairs. Its distribution is more tropical, from

Rhodesia to northern South West Africa and northern

Cape. It overlaps with H. petrensis but the com-

bination of narrow leaves (less than 5 mm wide) and

subglabrous, somewhat varnished stems, serves to

identify H. linearis.

H. petrensis, with villous stems and with leaves rarely

narrower than 5 mm, varies a good deal in leaf shape

from narrowly lanceolate to ovate-lanceolate, while

the pubescence of the leaves may be villous to shortly

canescent, or a mixture of the two. Occasional inter-

mediates are found, for example the two specimens

from the Waterberg Plateau in S.W. Africa might

be better placed in H. canescens, and further investi-

gation in this area is desirable. Occasionally the

stems and leaves are sparingly villous, approaching

the condition of H. linearis. Orthosiphon mossianus

was based on such a plant. Ashby, l.c., distinguished

it from H. petrensis on the presence of petioles, but

this varies as much as the degree of pubescence and

leaf shape, so that O. mossianus and O. varians

(ovate-lanceolate leaves) are best regarded as forms

of H. petrensis. O. holubii is in no way distinct from

typical H. petrensis.

The types of O. dinteri and O. engleri have not been

seen. Launert & Schreiber, l.c., include them in

H. petrensis and, judging from the descriptions, this

appears to be the correct decision.

On this basis, H. petrensis is distributed from

Angola and Rhodesia southwards to Windhoek in

S.W. Africa and to the northern and eastern Transvaal

lowveld, reaching as far south as Komatipoort.

Superficially it resembles H. bracteosa which has a

somewhat similar distribution in our area, but the

latter species has a conspicuous coma of white to

rose-purple bracts, the flowers are usually whitish, not

mauve to violet as in H. petrensis, and the corolla is

usually shorter.

28. Hemizygia bracteosa ( Benth .) Briq. in Annu.

Conserv. Jard. Bot. Geneve 2: 248 (1898); Ashby in

J. Bot. Lond. 73: 352 (1935); Launert & Schreiber in

Prodr. FI. S.W. Afr. 123: 12 (1969). Type: from

Senegal.

Ocimum bracteosum Benth., Lab. 14 (1832); in Hook. Icon. PI.

t. 455 (1842); in DC., Prodr. 12:41 (1848).

Orthosiphon schinzianus Briq. in Bot. Jahrb. 19: 173 (1894).

Type: Amboland, Schinz 45 (Z, holo.;K). O. bracteosus (Benth.)

Bak. in FI. Trop Afr. 5: 375 (1900); N.E. Br. in FI. Cap. 5,1:

248 (1910). O. rhodesianus S. Moore in J. Bot. Lond. 43: 50

(1905). Type: Rhodesia, Wankie, Eyles 132 (BM, holo.).

Hemizygia junodii Briq. in Annu. Conserv. Jard. Bot. Geneve

2: 249 (1898). Syntypes: Mozambique, Delagoa Bay, Junod

61; 235. — var. quintasii Briq., l.c. 249 (1898). Type Mozam-

bique, Delagoa Bay, Quintas s.n. H. hoepfneri Briq. in Bull.

Herb. Boiss. ser. 2,3 : 994 (1903). Type: S.W. Africa, Hereroland,

Hopfner 85. H. serrata Briq., l.c. 996 (1903). Syntypes: S.W.

Africa, Amboland, Rautanen s.n.; Wulfhorst 1.

Bouetia ocimoides A. Chev. in Mem. Soc. Bot. Fr. 2: No. 8d,

200 (1917). Type: from West Africa.

Herb, probably annual, 25-75 cm tall, sometimes

woody and branching below, forming a soft aromatic

bush; stems sparingly to densely pilose with long weak

multicellular hairs. Leaves sessile; blade narrowly

lanceolate to oblong-lanceolate 4-9 cm long, 8-24 mm

broad, hispidulous and darker above, sparingly to

densely canescent below; apex acute, base attenuate;

margin somewhat distantly serrate-dentate. Inflores-

cence paniculate or simple, lax, 12-30 cm long;

rhachis glandular-hispidulous; bracts large forming

a persistent apical coma, white to rose-purple,

5-10x4-8 mm; verticillasters 4-6-flowered, 1-2,5 cm

apart. Calyx 5 mm long at anthesis, glandular-villous,

enlarging in fruit; upper lip broadly ovate to sub-

rotund, purple, 2 mm long, decurrent; lower 4 teeth

deltoid subulate, up to 2 mm long, becoming bristle-

like. Corolla white or tinged with mauve (rarely

violet), 10-11 mm long, puberulous on the lips

otherwise glabrous; tube 7-8 mm long widening

gradually to 2,5-3 mm at the throat; upper lip a

minute appendage; lower lip concave, 3 mm long,

often deflexed. Stamens exserted by 5-6 mm, the lower

pair longer than the upper pair, exceeding the lower

lip of the corolla; upper pair attached below the

middle of the tube, filaments finely puberulous for

half or more of their length; lower pair attached at

the throat, filaments united to near the apex. Stigma

somewhat clavate, entire.

Widespread from Senegal and Tanzania southwards

to northern South West Africa, northern Botswana,

Mozambique and the eastern Transvaal Lowveld.

Found among rocks, in watercourses and in open

sandy places in relatively dry tropical woodland.

S.W. A. — Grid uncertain: Koakoveld, Merxmiiller d Giess

1968; Grootfontein, near Gross Huis, Schoenfelder S 572;

S 803; Okavango, Mpungu, Soini s.n.; Omaheke, Dinter 2317

(SAM). 1713 (Swartbooisdrif) : 53 km N. of Ohopoho (-DA),

Giess d Leippert 7573. 1714 (Ruacana Falls): near Ruacana

(-AC), De Winter d Giess 7097. 1719(Runtu): 48 km E. of

Runtu (-CD), De Winter 4053. 1720 (Sambio): near junction of

Amuramba Omatako and Okavango (-CD), De Winter d Wiss

4141. 1724 (Katima Mulilo): near Katima Mulilo (-AC),

Killick d Leistner 3078; 24 km S.E. of Katima Mulilo (-CB),

Killick d Leistner 3295. 1816 (Namutoni): 64 km S.E. of Ondan-

gua (-AB), De Winter d Giess 6951. 1817 (Tsintsabis): between

Tsintsabis and Kuringkuru (-DB?), Schoenfelder S 530. 1819

(Karakuwise); Cigarette (-DC?), Maguire 2377. 1821 (Andara):

near Andara (-AB), Merxmiiller d Giess 1968: near Bagoni

(-BA), De Winter d Wiss 4390. 1917 (Tsumeb): near Tsumeb

(-BA), Dinter 1713 (SAM); 3010 (SAM); 7499; near Otavi

(-CB), Dinter 5700; Hardy 2135. 1918 (Grootfontein): 24 km

N. of Nurugas (-BB), Basson 19; farm Oliewenhof (-CB),

Giess, Volk d Bleissner 6521. 1919 (Kanovlei): 40 km S. of

Kanovlei (-AD). Giess 9819. 1920 (Tsumkwe): (-DA), Giess,

Watt & Snyman 11072; Aha Mts. (-DB), Story 6388; Guatscha

Pan (-DC), Story 6231. 2017 (Waterberg): Waterberg Plateau

(-AC), Boss sub TRV 35002. 2119 (Epukiro): Epukiro Reserve,

16 km E. of Abdreh (-AB?), Giess 9779. 2215 (Trekkopje):

Gaub (-CB), Dinter 2411 (SAM); 2433 (SAM).

Botswana. — Grid uncertain: Ngamiland, Curson 286;

557; 560; Buerger 1085; Ngami, Van Son sub TRV 28921;

Matsaudi, Lambrecht 81. 1725 (Livingstone): Kasane (-CC),

Biegel d Russell 3686. 1923 (Maun): Maun (-CD), Lambrecht

57. 2023 (Kwebe Hills): Kwebe Hills (-CA), Mason d Boshoff

275. 2125 (Lothlekane): Orapa (-AD), Allen 98.

20

THE SOUTH AFRICAN SPECIES OF HEMIZYGIA (LAMIACEAE)

Transvaal. — 2231 (Pafuri): 8 km N.W. of Punda Milia

(-CA), Bruce 171; Wambia area (-CB), Van der Schijff 2957;

Schlieben 9311; Dzundweni Hill (-CC), Codd 4260. 2431

(Acornhoek): Acornhoek (-CA), Roberts sub TRV 26204;

8 km from Newington to Bushbuck Ridge (-CD), Buitendag 912;

Kruger National Park, Sand River (-DC), Van der Schijff 2192.

2531 (Komatipoort): near Shabin Kop (-AA), Acocks 16668;

14 km N. of Pretorius Kop (-AA), Codd 5198; Pretorius Kop

Camp (-AB), Van der Schijff 273; Faai River (-AB), Ihlenfeldt

2361.

In habit, ecology and distribution within our area,

H. bracteosa resembles the former species, H. petrensis,

but may be distinguished by the conspicuous coma

of large, whitish to rose-purple bracts, the usually

whitish corolla which is usually shorter than the

mauve to violet corolla of H. petrensis. The leaves are

canescent as in H. canescens but the stems are weakly

pilose, while the conspicuous bracts distinguish it

from the latter species. H. bracteosa is remarkably

uniform considering its wide distribution from

Senegal and Tanzania to Southern Africa.

UITTREKSEL

'n Oorsig oor die Suid-Afrikaanse species van

Hemizygia word gegee en 28 species word herken,

insluitende die volgende nuwe name; H. macrophylla

( Guerke ) Codd (=Syncolostemon macrophyllus

Guerke), H. pretoriae Guerke var. heterotricha Codd ,

H. cinerea Codd, H. incana Codd, H. modesta Codd, H.

parvifolia Codd, H. punctata Codd en H. ramosa Codd

INDEX

Bouetia A. Chev

ocimoides A. Chev

Hemizygia ( Benth .) Briq

albiflora ( N.E.Br .) Ashby

bolusii (N.E.Br.) Codd

bracteosa (Benth.) Briq

canescens (Guerke) Ashby . .

cinerea Codd

elliottii (Bak.) Ashby

floccosa Launert

foliosa S. Moore

gerrardii (N.E.Br.) Ashby ....

hoepfneri Briq

humilis (N.E.Br.) Ashby

incana Codd

junodii Briq

var. quintasii Briq

linearis (Benth.) Briq

macrophylla (Guerke) Codd .

modesta Codd

mossiana (Good) Ashby ....

obermeyerae Ashby

parvifolia Codd

persimilis (N.E.Br.) Ashby . .

petiolata Ashby

petrensis (Hiern) Ashby ....

pretoriae (Guerke) Ashby .. .

subsp. heterotricha Codd.. .

subsp. pretoriae

punctata Codd

ramosa Codd

rehmannii (Guerke) Ashby . .

rehmannii sensu Compton . .

rugosifolia Ashby

serrata Briq

stenophylla (Guerke) Ashby .

aff. stenophylla

subvelutina (Guerke) Ashby . .

teucriifolia (Hochst.) Briq. . .

thorncroftii (N.E.Br.) Ashby .

transvaalensis (Schltr.) Ashby

Nautochilus Brem

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Ocimum L

bracteosum Benth

Orthosiphon Benth

affinis N.E. Br

albiflorus N.E. Br

bolusii N.E. Br

bracteosus (Benth. )Bak

canescens Guerke

decipiens N.E. Br

dinteri Briq

elliottii Bak

engleri Perkins

foliosus (S. Moore) N.E. Br

galpiniana Briq

gerrardii N.E. Br

holubii N.E. Br

humilis N.E. Br

linearis Benth

macrophyllus (Guerke) N.E. Br. . .

messinensis Good

mossianus Good

muddii N.E. Br

natalensis Guerke

persimilis N.E. Br

petrensis Hiern

pretoriae Guerke

rehmannii Guerke

rhodesianus S. Moore

rogersii N.E. Br

schinzianus Briq

stenophyllus Guerke

subvelutina Guerke

teucriifolius (Hochst.) N.E. Br

var. galpinianus (Briq.) N.E. Br.

thorncroftii N.E. Br

transvaalensis Schltr

varians N.E. Br

wilmsii Guerke

woodii Guerke

Syncolostemon E. Mey. ex Benth. . .

macrophyllus Guerke

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Bothalia 12, 1: 21-27 (1976)

The genus Syncolostemon (Lamiaceae)

L. E. CODD*

ABSTRACT

The genus Syncolostemon E. Mey. ex Benth. is revised and its relationship with Hemizygia (Benth.) Briq.'

is discussed. A new species, S. comptonii Codd is described and the following new combinations are made:

S. parviflorus var. lanceolatus (Guerke) Codd (=S. lanceolatus Guerke) and S. latidens (N.E. Br.) Codd

( = Orthosiphon latidens N.E. Br.).

Resume

LE GENRE SYNCOLOSTEMON (. LAMIACEAE )

Le genre Syncolostemon E. Mey .ex Benth. est revise et sa relation avec Hemizygia (Benth.) Bri .esl discutee.

Une nouvelle espece, S. comptonii Codd est decrite et les combinaisons nouvelles suivantes sont faites: S. parvi-

florus var. lanceolatus ( Guerke ) Codd ( = S. lanceolatus Guerke) et S. latidens (N.E. Br.) Codd ( = Orthosiphon

latidens N.E. Br.).

INTRODUCTION

In his revision of the genus Hemizygia Briq. in J.

Bot. Lond. 73: 343 (1935), Ashby drew attention to

the floral differences between this genus and Ortho-

siphon Benth. but hardly touched on the genus

Syncolostemon E. Mey. ex Benth.

The corolla characters in Syncolostemon and

Hemizygia are essentially the same and lend no

support for their separation as distinct genera: the

tube is longer than the calyx, straight or slightly

decurved, widening about the middle towards the

truncate mouth; the upper corolla lip is usually very

small, obscurely 4-lobed, while the lower lip is

shallowly concave and longer than the upper lip.

The stamens are in two distinct pairs with the upper

(posticous) pair attached at or below the middle of

the corolla tube, with the filaments free and usually

pubescent towards their base; the lower (anticous)

pair are attached at the corolla throat with the fila-

ments united (united only near the base in H. persimilis

and somewhat variable in H. pretoriae).

The only character which shows some promise as a

basis for separating the two genera is the shape of

the calyx. In typical Syncolostemon the calyx is more

or less equally 5-toothed, while in Hemizygia the

upper calyx tooth is broadly ovate, often decurrent

on the calyx tube, and much larger than the lower

four which are usually subulate to almost spinescent.

There are, however, certain intermediates which are

discussed more fully below.

Syncolostemon E. Mey. ex Benth. in E. Mey.,

Comm. 230 (1837) was erected to accommodate

certain plants collected by Drege. Four species were

described which fall into two distinct groups:

Group 1 : S. parviflorus E. Mey. ex Benth. Calyx

and corolla small: Mature calyx less than 8 mm long,

campanulate, often becoming subrotund, setose in

the throat; corolla tube 6-10 mm long; inflorescence

lax; nutlets with a frill at the base.

Group 2: S. rotundifolius E. Mey. ex Benth. and

S. densiflorus Benth. (=S. ramulosus E. Mey. ex

Benth.). Calyx and corolla large and conspicuous:

mature calyx usually exceeding 10 mm in length,

cylindrical; corolla tube 14-30 mm long; inflorescence

dense; nutlets without a frill at the base.

To Group 1 the following may be added: S.

concinnus N.E. Br., S. argenteus N.E. Br., S. erioce-

phalus Verdoorn and the presently described S.

* Botanical Research Institute, Department of Agricultural

Technical Services, Private Bag X101, Pretoria.

comptonii Codd. The species S. macrophyllus Guerke,

although showing a distant relationship with S.

parviflorus, has a broadly ovate upper calyx tooth and

is transferred to Hemizygia.

Closely related to Group 2 are two species described

as Orthosiphon macranthus Guerke and O. latidens

N.E. Br. in which the upper calyx tooth is distinctly

larger than the lower four, but is elliptic to broadly

elliptic rather than broadly ovate. These two species

are therefore, somewhat intermediate between Syn-

colostemon and Hemizygia and Ashby transferred the

former to Syncolostemon and the latter to Hemizygia.

They are both so obviously related to S. rotundifolius

that it is logical to place all three in the same genus.

It is a matter of convenience to maintain both

genera and this may be done on the following basis:

Calyx 2-lipped, the upper lip broadly ovate to subrotund,

usually decurrent on the tube; lower lip of 4 subulate to

spinescent teeth Hemizygia

Calyx subequally 5-toothed, the upper tooth, if the largest,

elliptic to broadly elliptic, not decurrent; lower 4 teeth

deltoid to subulate Syncolostemon

Phillips, Gen. ed. 2: 651 (1951), lists S. densiflorus

as the lectotype species without giving reasons for

his choice. It will be noted that 5. densiflorus belongs

to Group 2, which tends to grade into Hemizygia.

Should this group be merged with Hemizygia, the

older name Syncolostemon would then take prece-

dence, with the result that some 30 names in Hemizygia

would have to be transferred to Syncolostemon; on

the other hand, Group 1 might be considered

sufficiently different to be maintained as a distinct

genus and would need a new name. It is therefore

considered that a better choice of lectotype would be

S. parviflorus E. Mey. ex Benth., belonging to Group 1

SYNCOLOSTEMON

Syncolostemon E. Mey. ex Benth. in E. Mey.,

Comm. 230 (1837); Benth. in DC., Prodr. 12: 53

(1848); Benth. & Hook, f., Gen. PI. 2,2: 1174 (1876);

Briq. in Pflanzenfam. 4,3a: 364 (1897); N.E. Br. in

FI. Cap. 5,1: 261 (1910); Phillips, Gen. ed. 2: 651

(1951); Ross, FI. Natal 306 (1972).

Lectotype proposed: S. parviflorus E. Mey. ex

Benth.

Soft shrubs or stems herbaceous, arising annually

from a perennial woody rootstock. Leaves opposite,

subsessile or shortly petiolate, variously pubescent

and gland-dotted, entire or toothed. Inflorescence

terminal, sometimes on short lateral shoots, paniculate

or simple, lax or dense; verticillasters 2-6-flowered

22

THE GENUS SYNCOLOSTEMON (LAMIACEAE)

Calyx variously pubescent and often setose in the

throat, subequally 5-toothed or the upper tooth

larger than the lower 4, elliptic to broadly elliptic, not

decurrent on the tube; lower 4 teeth deltoid to

subulate. Corolla bilabiate; tube longer than the

calyx, widening from about the middle to a truncate

mouth; upper lip small, obscurely 4-lobed; lower lip

concave, horizontal, longer than the upper lip.

Stamens 4, exserted, didynamous; upper pair affixed

about the middle or towards the base of the corolla

tube, filaments free, pubescent below; lower pair

Corolla tube 6-10 mm long (Group 1):

Pubescence on leaves of stellate hairs

Pubescence on leaves dense or sparse but not

attached at the corolla throat, filaments connate for

practically their entire length, glabrous. Style exserted,

minutely bilobed at the apex. Nutlets oblong in outline,

sometimes slightly frilled at the base.

Endemic in south-east Africa, extending from the

Transkei to the Transvaal; 9 species recognized.

Briquet, l.c., divided the species into two sections,

corresponding to Groups 1 and 2 above, which he

designated as Sect. Micranthi and Sect. Macranthi,

respectively.

1.5. concinnus

Leaves greenish, sparsely to densely pubescent but not sericeous:

Inflorescence lax, verticillasters up to 2 cm apart ; bracts ovate-lanceolate, not chartaceous :

Leaves obovate or elliptic to lanceolate, up to 3 times as long as broad .... 2a. 5. parviflorus

var. parviflorus

Leaves narrowly lanceolate to linear-lanceolate, 4-8 times as long as broad

2b. S. parviflorus var. lanceolatus

Inflorescence dense, verticillasters 2-3 mm apart; bracts broadly ovate, chartaceous

4. S. comptonii

Leaves grey, densely sericeous:

Leaves 12-25x2-8 mm, flat; inflorescence terminal, lax 3. S. argenteus

Leaves 4-10 x 1 ,5-3 mm, margin revolute; inflorescence terminal or on short lateral

shoots, dense, densely villous 5. S. eriocephalus

Corolla tube 14-30 mm long (Group 2):

Calyx teeth subequal:

Verticillasters usually 6-flowered ; calyx teeth linear-subulate, 3-5 mm long .... 6. 5. densiflorus

Verticillasters 2-flowered; calyx teeth narrowly deltoid, 1,5-2 mm long. ..7. S. rotundifolius

Calyx with the upper tooth elliptic to broadly elliptic, larger than the lower 4:

Corolla tube 2-3 cm long; rhachis glandular-puberulous 8. S. macranthus

Corolla tube 1,8-2 cm long; rhachis hispid 9. S. latidens

1. Syncolostemon concinnus N.E.Br. in FI.

Cap. 5,1: 264 (1910). Type: Transvaal, Elandspruit

Mts. (Steenkampsberg), Schlechter 3891 (K, holo.;

PRE).

Shrublet 60 cm tall; stems several, annual, slender,

sparingly branched, arising from a perennial woody

rootstock, beset with tufts of leaves along the stem,

hispidulous. Leaves subsessile obovate-oblong to

narrowly elliptic, 10-18 mm long, 2-8 mm broad,

with minute stellately branched hairs on both surfaces,

freely gland-dotted; apex rounded, base cuneate;

margin often with a few teeth near the apex. Inflores-

cence a lax panicle 12-20 cm long; rhachis glandular-

puberulous often with stellate hairs; bracts very small,

2-3 mm long, caducous; verticillasters 2-flowered,

up to 2 cm apart. Calyx 5-6 mm long, campanulate,

becoming subrotund with a narrowed mouth, glan-

dular-hispid, often with stellate hairs, setose in the

mouth; 5 teeth subequal, deltoid-subulate, 2,5 mm

long. Corolla white, 12—13 mm long; tube 8-9 mm

long; upper lip small, erect; lower lip horizontal,

3 mm long. Stamens well exserted; upper pair attached

near the base of the corolla tube, filaments glabrous

below.

Found in mountain grassland, usually among rocks,

in eastern and south-eastern Transvaal, southern

Swaziland and north-eastern Orange Free State;

main flowering season is February-March.

Transvaal. — 2530 (Lydenburg): Steenkampsberg (-AA),

Schlechter 3891 ; Codd 9847; Belfast (-CA), Leendertz sub TRV

7891. 2630 (Carolina): S.E. of Ermelo (-CA), Codd 10207;

Iswepe (-DC), Sidey 1617.

O.F.S. — 2729 (Volksrust): Top of Normandien Pass (-DC),

Acocks 23803.

Swaziland. — 2631 (Mbabane): 8 km W. of Mankaiana

(-CA), Compton 28664; Hlatikulu (-CD), Compton 28757.

With its slender stems and small white flowers,

S. concinnus is an inconspicuous plant, closely related

to the next species S. parviflorus. It differs in having

stellate (branched) hairs, often minute, on the leaves

and sometimes on other parts as well. The two

specimens from Swaziland, Compton 28664 and 28757,

differ in having very dense stellate pubescence on the

leaves, rhachis and calyx. This is the most northerly

member of the genus, extending to the Lydenburg

District.

2. Syncolostemon parviflorus E. Mey. ex Benth.

in E. Mey. Comm. 231 (1837). Lectotype: near

Umsikaba River, Drege (K, lecto.).

Shrublet 40-100 cm tall: stems few to several,

slender, sparingly branched, usually arising annually

from a woody rootstock, beset with tufts of leaves

along the stem. Leaves subsessile, greenish or drying

blackish, elliptic-obovate to lanceolate-elliptic or

linear, 12-32 mm long, 2-12 mm broad, hispidulous

to fairly densely appressed pubescent; apex rounded

to acute, base cuneate; margin with occasional teeth

near the apex on the larger leaves. Inflorescence a lax

panicle, 12—25 cm long; rhachis hispidulous; bracts

very small, ovate-lanceolate, 3-4 mm long, caducous;

verticillasters 2-flowered, up to 2 cm apart. Calyx

5-6 mm long, campanulate, becoming subrotund

often with a narrowed mouth, glandular-hispidulous,

setose in the mouth; 5 teeth subequal, deltoid-subulate,

2 mm long. Corolla white or flushed with pink,

rarely reddish-pink, 10-11 mm long; tube 6-7 mm

long; upper lip small, erect; lower lip horizontal,

3 mm long. Stamens well exserted, upper pair attached

near the base of the corolla tube, filaments with a

few hairs.

Found from the Transkei to Barberton District in

the Transvaal, in dense grassland, often among rocks,

from near sea level in Natal in 1 800 m altitude in

the Transvaal; flowering is mainly from January to

March.

L. E. CODD

23

Two varieties are recognized; for key to varieties,

see key to species.

(a) var. parviflorus.

S. parviflorus E. Mey. ex Benth. in E. Mey., Comm.

231 (1837); Drege, Zwei Doc. 151 (1843); Benth. in

DC., Prodr. 12: 54 (1848): N.E. Br. in FI. Cap.

5,1: 263 (1910); Compton, FI. Swaz. 67 (1966); Ross,

FI. Natal 306 (1972) (as “parvifolius”). Lectotype:

near Umsikaba River, Drege (K, holo.). —var.

dissitiflorus (Benth.) N.E. Br., l.c. 264 (1910). S.

dissitiflorus Benth. in DC., Prodr. 12: 54 (1848).

Type: Port Natal, Drege (K, holo.). S. lanceolatus

sensu Compton, FI. Swaz. 67 (1966).

Leaves obovate to oblanceolate or elliptic, 12-26x

4-12 mm.

Distribution and ecology as for the species.

Transvaal. — 2531 (Komatipoort): between Barberton and

Havelock (-CC), Codd 9530; Hilliard & Burn 3664.

Swaziland.- — 2531 (Komatipoort): Piggs Peak (-CC),

Miller 5208; 7252. 2631 (Mbabane): Forbes Reef (-AA),

Compton 25568; 31967; Mbabane (-AC), Codd 9510; Compton

24828; 25333; 25568; 26762; 27412; 30012; Schlieben 9573;

near Ukutu Forest (-AC), Meeuse 10140.

Natal. — 2730 (Vryheid): Utrecht, farm Naauwhoek (-CB),

Devenish 421. 2731 (Louwsburg): 11 km W. of Ngome (-DC),

Codd 9476. 2830 (Dundee): Kranskop (-DD), Codd 9664.

2831 (Nkandla): Nhlazatshe Mt. (-AA), Hilliard & Burn 3336.

2930 (Pietermaritzburg): The Dargle (-AC), Letty 265; Umlazi

(-CD), Medley Wood 9652: Inanda (-DB), Strey 5168; Krantz-

kloof (-DD), Medley Wood 12015. 2931 (Stanger): Groutville

(-AD), Moll 2564; Durban (-CC), Krauss 145. 3030 (Port

Shepstone): Mtwalume (-BC), Hilliard <£ Burn 3396; Nicholson

1135; Ellermere (-BC), Rudatis 1384; Umgayi (-BC), Ward

5492. 3130 (Port Edward): near Port Edward (-AA), Nicholson

992.

Cape. — 3129 (Port St. Johns): near Lusikisiki (-BC), Acocks

13248; Magwa Falls (-BC), Galpin 10961.

Superficially resembles S. concirtnus but hairs on

leaves, rhachis etc. simple, not stellate. It grades into

var. lanceolatus, listed below, but is distinguished by

the broader leaves. In the great majority of specimens

the leaves and stems are shortly hispid but in Codd

9576 and 9664 from central Natal they are distinctly

villous.

(b) var. lanceolatus ( Guerke ) Codd, stat. nov.

Syncolostemon lanceolatus Guerke in Bot. Jahrb.

26: 77 (1898); N.E. Br. in FI. Cap. 5,1: 262 (1910);

Ross, FI. Natal 306 (1972). Lectotype: East

Griqualand, Mt. Malowe, Tyson in Herb. Norm.

Austr. Afr. 1294 (K, lecto.; PRE). — var. grandiflorus

N.E. Br., l.c. 262 (1910). Type: Natal, Enon, Wood

1882 (K, holo.). — var. cooperi (Briq.) N.E. Br., l.c.

262 (1910). S. cooperi Briq. in Bull Herb. Boiss. ser.

2,3: 979 (1903). Syntypes: Natal, Cooper 1151;

2895.

Leaves narrowly lanceolate to linear-lanceolate,

often appressed-pubescent on both sides.

Recorded from East Griqualand and the central

Natal midlands. N. E. Brown records Cooper 2895 as

having been collected in the Orange Free State, but

this seems unlikely and requires confirmation.

Natal. — 2930 (Pietermaritzburg): Greytown (-BA), Medley

Wood 9909; Galpin 14691 ; near Harburg (-BC), Marais 801;

New Hanover (-BC), Wells 1882; Noodsberg (-BD), Moll 1488;

Pietermaritzburg (-CB), Medley Wood 11121; Moss 7290;

Byrne (-CC), Galpin 14691; 25 km S.W. of Pietermaritzburg

(-CD), Dyer 4707; Tala, 8 km S.W. of Thornville (-CD),

Moll & Morris 636.

Cape. — 3029 (Kokstad): near Clydesdale (-BD), Tyson 1762;

2770; Schlechter 6616.

Grades into var. parviflorus and so varietal status

appears appropriate. There is also an indication of

introgression with S. argenteus, especially where the

two tend to meet in Pietermaritzburg district.

S. argenteus is a more robust species with a semi-

coastal distribution and with markedly sericeous

leaves.

3. Syncolostemon argenteus N.E Br. in FI. Cap.

5,1: 263 (1910); Ross, FI. Natal 306 (1972). Type:

Natal, near Inyezaan, Medley Wood 3875 (K, holo.).

Herb or soft shrublet 60-130 cm tall; stems solitary

or few from the base, slender, sparingly branched,

sericeous, beset with tufts of leaves along the stem.

Leaves subsessile, linear-lanceolate to elliptic-obovate,

12-25 mm long, 2-8 mm broad, densely sericeous,

entire. Inflorescence a fairly lax panicle, 9-25 cm

long; rhachis hispid to sericeous; bracts very small,

2,5-3 mm x 1,5 mm, caducous; verticillasters 2-

flowered, up to 1,5 cm apart. Calyx 5-6 mm long,

campanulate, becoming subrotund, glandular-hispid,

villous in the mouth; 5 teeth subequal, deltoid-

subulate, 1,5 mm long. Corolla white to pinkish,

10-12 mm long; tube 8 mm long; upper lip small,

erect; lower lip horizontal 3 mm long. Stamens well

exserted, upper pair attached near the middle of the

tube, filaments with a few hairs.

Found in dense grassland, often adjoining forest,

in central Natal midlands and semi-coastal areas at

altitudes of 300 to 1 000 m.

Natal. — 2831 (Nkandla): Nhlazatshe Mt. (-AA), Hilliard &

Burn 3326; Mpembeni, near Hlabisa (-BB), Ward 3078;

Nkandla Forest (-CA), Wells 2496; De Winter 8262; Ntumeni

(-CD), Medley Wood 9361 ; Eshowe (-CD), Strey 4592; Ngoya

(-DC), Medley Wood 10359; Mtunzini (-DC), Mogg 4343;

4422; 5808. 2930 (Pietermaritzburg): 16 km S. of Kranskop

(-BB), Moll 1664; near Pietermaritzburg (-CB), Acocks 13449;

Dohse & Lindahl 93; Pateni Estates (-CC), Nicholson 535; Mid

Illovo (-DC), Sim 19863. 3030 (Port Shepstone): Umgayi

(-BC), Ward 5495; Isezela (-BC), Strey 6475.

Allied to S. parviflorus but is somewhat more robust

with grey, silky-pubescent leaves. Some specimens

with narrow leaves are not always distinguishable with

certainty from S. parviflorus \ ar. lanceolatus, especially

in the Maritzburg area where the two overlap,

suggesting introgression between the two.

4. Syncolostemon comptonii Codd, sp. nov., a

S. parvifloro E. Mey. ex Benth. inflorescentia com-

pacta, verticillastris 2-3 mm separatis differt.

Suffrutex 160 cm altus; caules graciles, erecti,

breviter antrorsi-pilosi. Folia subsessilia, oblanceolata

vel anguste elliptica, 2-3,5 cm longa, 3-6 mm lata,

parce hispidula, copiose glanduloso-punctata, apice

acuto, basi anguste cuneata, margine integro vel apice

minute dentato. Inflorescentia compacta, paniculata,

5-8 cm longa, 2-2,5 cm lata; rhachis glanduloso-

hispidula; bracteae chartaceae, late ovatae, 2,5 X

2,5 mm, caducae, margine villoso; verticillastri

2-floribus, 2-3 mm separati; pedicelli 1 mm longi,

villosi. Calyx per anthesin 6 mm longus, hispidus,

glanduloso-punctatus, fauce setosa; dentes 5, sub-

aequales, deltoideo-subulati, 1,5 mm longi. Corolla

alba, 9-10 mm longa, glabra vel extus labiis puberulis;

tubus 6 mm longus, apicem versus ampliatus, ore

2 mm lato; labium posticum 1 mm longum; anticum

concavum, 3 mm longum. Stamina 9 mm exserta;

postica circa medium tubi corollae inserta, filamentis

libris basin versus parce pubescentibus; antica fauce

corollae, inserta, filamentis ad apicem connatis.

Stylus 8 mm exsertus, apice breviter bifido.

Type: Swaziland, 2631 (Mbabane), near Komati

Bridge (-AA), Compton 28839 (PRE, holo.).

Soft shrub 160 cm tall; stems slender, erect, shortly

antrorse-pilose. Leaves subsessile, oblanceolate to

24

THE GENUS SYNCOLOSTEMON (LAMIACEAE)

narrowly elliptic, 2-3,5 cm long, 3-6 mm broad,

sparingly hispid, freely gland-dotted; apex acute, base

narrowly cuneate; margin entire or with the apex

minutely dentate. Inflorescence compact, paniculate,

5-8 cm long, 2-2,5 cm wide; rhachis glandular-

hispidulous; bracts chartaceous, broadly ovate,

2 , 5 X 2 , 5 mm, caducous, margin villous ; verticillasters

2-flowered, 2-3 mm apart; pedicels 1 mm long,

villous. Calyx at flowering 6 mm long, hispid, gland-

dotted, throat setose; teeth 5, subequal, deltoid-

subulate, 1 ,5 mm long. Corolla white, 9-10 mm long,

glabrous with the outer surface of the lobes

puberulous; tube 6 mm long, widening towards the

apex, mouth 2 mm wide; posticous lobe 1 mm long;

anticous lobe concave, 3 mm long. Stamens exserted

by 9 mm ; posticous stamens inserted about the middle

of the corolla tube, filaments free, sparingly pubescent

towards the base; anticous stamens inserted at the

throat of the corolla, filaments united to the apex.

Style exserted by 8 mm, apex shortly bifid. Fig. 1.

it' y ■ k •

Do. ^

Fig. 1. — Syncolostemon comptonii ( Compton 28839, PRE, holotype)

L. E. CODD

25

Known only from the single gathering cited above,

from Komati Bridge on the road from Mbabane to

Piggs Peak.

Although closely related to 5. parviflorus E. Mey.

ex Benth., the specimen appears to be sufficiently

distinct to be separated as a species. It is taller (160

cm), the inflorescence is compact with verticillasters

2-3 mm apart, and the bracts are chartaceous,

broadly ovate, as broad as long, with a fringe of woolly

hairs. In S. parviflorus the stems are 40-100 cm tall,

the inflorescence is lax with verticillasters up to 2 cm

apart, and the bracts are ovate-lanceolate, without a

fringe of woolly hairs.

5. Syncolostemon eriocephalus Verdoorn in Kew

Bull. 1937: 447 (1937). Type: Transvaal, Pilgrim’s

Rest, Morisse 51.

Shrub 60-200 cm tall, much branched; branches

beset with closely placed tufts of leaves along the

stem; branchlets densely villous; Leaves sessile, small,

linear or narrowly oblong, 4-10 mm long, 1 ,5-3 mm

broad, densely silvery sericeous on both surfaces;

apex rounded, base shortly cuneate, margin entire.

Inflorescence terminal or on short lateral shoots,

paniculate or simple, dense, 2-5 cm long; rhachis

densely villous; bracts ovate, acuminate, 4-6 mm long,

densely villous, caducous; verticillasters 2-flowered,

1-2 mm apart; pedicels 1-1,5 mm long. Calyx 4 mm

long, thickly covered with white to pale yellowish

woolly hairs, villous in the mouth; 5 teeth subequal,

deltoid-acuminate. Corolla cream, yellow or brownish-

yellow, 7-9 mm long; tube 5-6 mm long; upper lip

very small; lower lip concave, horizontal, 2 mm long.

Stamens exserted by 3 mm; upper pair attached

about the middle of the corolla tube, filaments free,

pubescent near the base.

Found in shallow sandy soil among quartzite rocks

along the Drakensberg escarpment from Pilgrim’s

Rest to near The Downs, at altitudes of 1 400 to

2 000 m. Collected in flower from July to February.

Transvaal. — 2430 (Pilgrim’s Rest): 8 km S.E. of The Downs

(-AA), McNeil s.n.; Mariepskop (-DB), Keet 1111; Van der

Schijff 4838; 5587; 6760; 7328; Killick & Strey 2378; farm

Belvedere (-DB), Codd 10314; Lisbon River (-DD), Liebenberg

3551 ; Galpin 14601; Jordaan 98; 18 km N. of Graskop, Codd &

de Winter 3349; near Pilgrim’s Rest (-DD), Morisse 51 ; Rauh &

Schlieben 9661.

A very distinctive species because of its shrubby

habit, small grey leaves, dense small inflorescences

nearly obscured by woolly hairs and small cream to

yellowish-brown flowers.

6. Syncolostemon densiflorus Benth. in E. Mey.,

Comm. 230 (1837); Drege, Zwei Doc. 144, 149

(1843); Hochst. in Flora 67 (1845): Benth. in DC.,

Prodr. 12: 54 (1848); N.E. Br. in FI. Cap. 5,1: 265

(1910); Codd in Flow. PI. Afr. 32: t. 1252 (1957);

Ross, FI. Natal 306 (1972). Lectotype: Cape,

between Umzimkulu River and Magwa Falls,

Drege 4744 c (K, lecto.).

S. ramulosus E. Mey. ex Benth. in E. Mey., Comm. 231

(1837); Drege, Zwei Doc. 148, 152 (1843); Benth. in DC.,

Prodr. 12: 54 (1848); N.E. Br. in FI. Cap. 5,1: 264 (1910).

Syntypes: near Morley, Drege; near Umsikaba River, Drege.

Shrub 1-2,2 m tall, sparingly branched; branches

shortly white tomentose. Leaves petiolate; petiole

1-6 mm long; blade ovate or broadly elliptic to

orbicular, 5-15 mm long, 4-10 mm broad, scabrid-

hispidulous to subglabrous, gland-dotted; apex obtuse

to acute, base cuneate; margin subentire or toothed

above the middle. Inflorescence a dense terminal

panicle 5-16 cm long, 4-6,5 cm in diameter; rhachis

finely appressed tomentulose; bracts caducous,

broadly ovate to orbicular, 3-5 x 3-4 mm, cuspidate,

margin ciliate; verticillasters 4-6-flowered, 2-3 mm

apart, pedicels 1 mm long. Calyx 10 mm long, cylin-

drical, sparingly hispidulous, not villous in the mouth;

5 teeth subequal, deltoid-subulate, 2,5-5 mm long,

the uppermost tooth often slightly shorter than the

rest, suberect. Corolla crimson, pink or rarely whitish,

1 8—23 mm long; tube 15-21 mm long, gradually

widening to 5-6 mm wide at the mouth, glabrous,

with lips finely puberulous; upper lip short; lower lip

concave, 3-4 mm long, usually deflexed at maturity.

Stamens exserted by 10-12 mm, often coiled; upper

pair inserted below the middle of the corolla tube,

filaments minutely puberulous near the base.

Distributed from Keiskamma Hoek in the eastern

Cape Province to about Nongoma in Natal, in dense

grassland often adjoining forest, at altitudes up to

1 000 m. It has been collected in flower between

November and July, but the main flowering time is

from February to May.

Natal. — 2731 (Louwsburg): near Nongoma (-DC), Ward

4066. 2831 (Nkandla): Nkandla Forest (-CA), Edwards 1397;

between Nkandla and Eshowe (-CC), Strey 4161 ; Eshowe (-CD)

Galpin 13553; Rogers 24889; McClean 982; Johnson 1428;

Umlalazi (-DD), Wylie sub Wood 10340; Mtunzini (-DD),

Huntley 199. 2930 (Pietermaritzburg): Pietermaritzburg (-CB),

Sidey 3541; between Hella-Hella and Richmond (-CC),

Edwards 3112; near Richmond (-CD), Acocks 13778; Bayliss

2195; Ismont (-DC), Strey 8369. 2931 (Stanger): Doornkop

Estates (-AA), Pentz & Acocks 10426; St. Philomina Mission

(-AA), Moll 1619. 3029 (Kokstad): Harding (-DB), Bayliss

2227. 3030 (Port Shepstone): Dumisa (-AD), Rudatis 232;

Umgayi (-AD), Ward 5487; near Port Shepstone (-CB),

Dimmock-Brown 391; Letty 237. 3130 (Port Edward): Port

Edward (-AA), Strey 6407.

Cape. — 3929 (Kokstad): Weza (-DA), Strey 11141;

Clydesdale (-BD), Tyson 1761 ; 2545; S. of Umzimkulu (-BD),

Acocks 12250; Story 659; Codd 8565; Tabankulu Mt. (-CB),

Hilliard & Burtt 3519. 3129 (Port St. Johns): near Lusikisiki

(-BC), Galpin s.n.; Comins 1920; Port. St. Johns (-DA), Bolus

10256; Hilliard & Burtt 3531. 3130 (Port Edward): Bizana

District (-AA), Codd 9340. 3227 (Stutterheim): Keiskamma Hoek

(-CA), Killick 885; Wells 3096; Kologha Forest (-CB), Theron

2139; Stutterheim (-CB), Wehrmeyer 10; Pirie (-CC), Sim

19586; Galpin 3260; Taylor 1756; near Komga (-DB), Flanagan

1890; Codd 9241; Nahoon Mouth (-DD), Galpin 7743. 3228

(Butterworth): near Manubi (-BC), Wells 3609; Plowes 2418;

Mauve 4879; Kentani (-CB), Pegler 386. 3327 (Peddie): East

London (-BB), Comins 1563.

The species may be distinguished by its subulate

calyx teeth 3-5 mm long, the upper tooth being often

shorter than the lower four. S. rotundifolius E. Mey

ex Benth. has shorter, more deltoid calyx teeth, and

the verticillasters are 2-flowered.

The type of S. ramulosus E. Mey. ex Benth. has

teeth somewhat intermediate between 5". densiflorus

and S. rodundifolius and, as it has 4-6-flowered

verticillasters, it is included in the former. No modern

material exactly matching it has been seen.

With its dense inflorescences of pink to reddish

flowers, 5. densiflorus makes an attractive garden

subject but attempts to cultivate it have not been

successful in the Transvaal.

7. Syncolostemon rotundifolius E. Mey. ex Benth.

in E. Mey., Comm. 231 (1837); Drege, Zwei Doc. 153,

155 (1843); Benth. in DC., Prodr. 12: 53 (1848);

N.E. Br. in FI. Cap. 5,1: 265 (1910); Ross, FI. Natal

306 (1972). Lectotype: Cape, between Umtentu and

Umsikaba Rivers, Drege 4743a (K, lecto.; PRE).

Soft shrub 60 cm-2 m tall, sparingly branched;

branches densely whitish appressed tomentulose.

Leaves petiolate; petiole 2-5 mm long; blade broadly

elliptic or broadly obovate to subrotund, 10-25 mm

long, 6-18 mm broad, shortly tomentulose and

usually gland-dotted; apex rounded, base cuneate to

obtuse; margin entire or faintly crenate-dentate above

26

THE GENUS SYNCOLOSTEMON (LAMIACEAE)

the middle. Inflorescence a fairly dense panicle, rarely

simple, 5-8 cm long, 5-6 cm in diameter; rhachis

densely appressed tomentulose; bracts caducous,

concave, broadly ovate, apiculate, 3-5 mm long,

3-4 mm broad, finely tomentulose; verticillasters

2-flowered, 3-5 mm apart, pedicels 1 mm long.

Calyx 9-10 mm long, cylindrical, finely tomentulose,

not villous in the mouth; 5 teeth subequal, narrowly

deltoid, 1,5-2 mm long, the uppermost tooth often

slightly shorter than the rest, suberect. Corolla mauve,

pink or magenta-pink, 23-27 mm long; tube 20-23 mm

long, gradually widening to 5-6 mm at the mouth,

glabrous; upper lip short; lower lip concave, 3-5 mm

long, usually reflexed at maturity. Stamens exserted by

12-15 mm, often coiled; upper pair inserted below the

middle of the corolla tube, filaments pubescent at the

base.

Found in grassland and scrub on rocky slopes from

about Port St. Johns to Port Shepstone, usually not

far from the sea at altitudes up to 400 m. Main

flowering season between December and May.

Natal. — 3030 (Port Shepstone): near Port Shepstone (-CB),

Strey 5798; Sidev 3542; Oribi Gorge (-CB), Burtt 2994; near

Paddock (-CC), McClecm 287; Hilliard 3975, Sidey 3870;

Uvongo (-CD), Whellen 1058; Munster (-CD), Gemmell s.n.;

Shelly Beach (-CD), Strey 8438. 3130 (Port Edward): near

Port Edward (-AA), Pole Evans s.n.; Acocks 13331 ; Bayliss 551 ;

Beacon Hill (-AA), Strey 6526; 6885.

Cape. — 3129 (Port St. Johns): Egossa (-BC), Sim 2521;

Magwa Falls (-BC), Codd 9321 ; 20 km S. of Lusikisiki (-BC),

Lewis sub SAM 68185; between Umzimkulu and Umtentu

Rivers (-BD?), Drege 4743a; Mkambati (-BD), Strey 8679;

Port St. Johns (-DA), Mogg s.n. 3130 (Port Edward): near

Umtamvuna Mouth (-AA), Pole Evans 4758; Acocks 10918;

Umtamvuna Waterfall (-AA), Strey 4473; Umzamba River

mouth (-AA), Lewis sub SAM 68184; Mtentu Location (-AA),

Strey 8638.

Related to S. densiflorus Benth., but the calyx teeth

are shorter and more deltoid, the verticillasters are

2-flowered, and the stems, leaves and calyx have a

short, soft appressed tomentum, as distinct from the

scabrid, hispidulous pubescence of S. densiflorus.

There is also a close resemblance with S. macranthus

(Guerke) Ashby of the Natal Drakensberg area, but

the latter has usually 4-6-flowered verticillasters, while

the upper calyx tooth is obovate-elliptic and distinctly

larger than the other four. The leaves of 5. macranthus

also tend to be larger and more acute at the apex,

with a short scabrid pubescence.

8. Syncolostemon macranthus ( Guerke ) Ashby in

J. Bot. Lond. 73: 357 (1935); Ross, FI. Natal 306

(1972). Lectotype; Natal, Van Reenens Pass, Medley

Wood 3573, in NH 949 (K, lecto.).

Orthosiphon macranthus Guerke in Bot. Jahrb. 26: 84 (1898);

N.E. Br. in FI. Cap. 5,1: 242 (1910).

Hemizygia cooperi Briq. in Bull. Herb. Boiss. ser. 2,3: 992

(1903). Type: “Orange Free State”, Cooper 1015.

Shrub 1-2,5 m tall, much branched; branches finely

hispidulous. Leaves petiolate; petiole 2-8 mm long;

blade ovate to ovate-lanceolate, 2-4,5 cm long,

1,2-2 cm broad, scabrid-hispidulous, gland-dotted;

apex acute to obtuse, base obtuse to cuneate; margin

obscurely crenate-dentate. Inflorescence simple or

usually branched, 8-18 cm long, fairly dense to lax;

rhachis glandular-puberulous; bracts caducous,

broadly ovate, acute, 8-10x5-6 mm, puberulous;

verticillasters 4-6 (rarely 2)-flowered, 4-18 mm apart;

pedicels 1 mm long. Calyx 9-10 mm long, cylindrical,

densely glandular-puberulous, not villous in the

mouth; upper tooth the largest, obovate-elliptic,

2-2,5 mm long, not decurrent on the tube; lower

4 teeth narrowly lanceolate, acuminate to subulate,

1 ,5-2 mm long. Corolla pink to pale mauve or purple,

25-30 mm long; tube 20-25 mm long, gradually

widening to 5-6 mm wide at the mouth, pubescent;

upper lip short; lower lip concave, 4-5 mm long,

usually deflexed at maturity. Stamens exserted by

10-12 mm, often coiled; upper pair inserted about

10 mm from the mouth of the tube, filaments glabrous.

Recorded from a restricted area of the Drakensberg

between Cathedral Peak and Van Reenens Pass,

where it is locally frequent along streams and forest

margins at altitudes of 1 600-2 200 m. Although

Cooper 1015 (K) is said to have been collected in the

“Orange River Colony”, this requires confirmation

as all modern gatherings are from the Natal side of

the border. Flowering mainly from January to May.

Natal. — 2828 (Bethlehem): Royal Natal National Park

(-DB), Trauseld 255. 2829 (Harrismith): Van Reenens Pass

(-AD), Medley Wood 12106; Schlechter 6912; Phillips s.n.;

Taylor 2013; Codd 8516; Oliviershoek Pass (-CA), Schweickerdt

sub TRV 31653; Acocks & Hafstrom 1340; Acocks 11207;

Bruce 405; Repton 1019; Codd 10533; Strey 9505; Umlambonja

Valley (-CC), Marriot s.n.; Saddle area (-CC), Edwards 2130;

Cathedral Peak Forest Station (-CC), Killick 1074; 1223;

1251; White 10586.

Differs from 5". rotundifolius in the calyx having

the upper tooth larger than the lower four, the

larger, more acute leaves and the fine scabrid pubes-

cence of stems and leaves.

5. latidens (N.E. Br.) Codd differs from it in the

leaves having a fine appressed tomentum and the

calyx hispid with the lower four calyx teeth deltoid-

lanceolate, not acuminate as in S. macranthus. The

two are closely related but the distribution is different,

S. latidens occurring in a restricted area near

Kranskop.

Briquet in Bull. Herb. Boiss. ser. 2,3: 993 (1903)

considered that H. cooperi Briq. occupied an isolated

position in Hemizygia and made it the type of a new

section, Sect. Cooperozygia Briq. H. cooperi is con-

specific with S. macranthus, which is so obviously

allied to S. rotundifolius that Ashby is considered

justified in placing it in Syncolostemon.

9. Syncolostemon latidens (N.E. Br.) Codd, comb.

nov.

Orthosiphon latidens N.E. Br. in FI. Cap. 5,1: 242 (1910).

Type: Natal, Umvoti District, Gerrard 1233 (K, holo.).

Hemizygia latidens (N.E. Br.) Ashby in J. Bot. Lond. 73 : 348

(1935); Ross, FI. Natal 306 (1972).

Soft shrub 1-1,5 m tall, branching; branches

hispidulous. Leaves petiolate; petiole 3-10 mm long,

tomentulose; blade ovate to broadly ovate, 3-5 cm

long, 2-3 , 5 cm broad, drying dark brown, tomentulose

on both surfaces, especially on the nerves; apex

obtuse to acute, base obtuse to truncate; margin

crenate-dentate. Inflorescence a fairly dense terminal

panicle, 10-20 cm long; rhachis hispid; bracts

caducous, broadly ovate, acute, 5 mm long, sparingly

pubescent; verticillasters 6-flowered, 4-8 mm apart;

pedicels 1 mm long, hispid. Calyx 10-11 mm long,

cylindrical, glandular-hispid, purple, not villous in

the mouth; upper tooth the largest, broadly obovate,

3-3,5 mm long, rounded at the apex, not decurrent;

lower 4 teeth lanceolate-deltoid, flat, tapering

gradually, 2,5 mm long. Corolla mauve-pink to deep

pink, 22-25 mm long; tube 20-22 mm long, widening

beyond the middle to 5-6 mm wide at the mouth,

finely pubescent, particularly on the lobes; upper lip

short; lower lip concave 3-5 mm long, often deflexed

at maturity. Stamens exserted by 10-12 mm; upper

pair inserted below the middle of the corolla tube,

puberulous near the base.

Known only from a restricted area near Kranskop

in central Natal, growing in and near the forest

margin. Flowering is mainly from March to May.

L. E. CODD

27

Natal. — Grid uncertain: Umvoti District, Gerrard 1233

(K); “ Greytown”, Wylie s.n.; Kranskop District, Olifants Hoek,

Strey 4248. 2930 (Pietermaritzburg): S.E. of Kranskop (-BB),

Dyer 4353; Codd 10193.

Closely related to S. macranlhus but differs in the

stro nger pubescence, the broader lower calyx teeth

and the somewhat shorter corolla. The leaves tend to

dry a dark brownish colour, which is not the case

with S. macranthus. These two species are somewhat

intermediate between Syncolostemon and Hemizygia

but the former genus is preferred because the upper

calyx tooth, although broad, is not decurrent on the

tube and the lower calyx teeth are not markedly

subulate or spine-like. They are also more closely

allied to S. rotundifolius and S. densiflorus than they

are to any Hemizygia species.

The four species are very showy and should make

good horticultural subjects. Unfortunately initial

attempts to cultivate them have not been successful

but they are worth persisting with until the difficulties

are overcome.

EXCLUDED SPECIES

Syncolostemon macrophyllns Guerke in Bull. Herb.

Boiss. 6: 555 (1898) is placed as Hemizygia macro-

phyllus ( Guerke ) Codd , comb. nov. (see p. 3).

UITTREKSEL

Die genus Syncolostemon E. Mey. ex Benth. word

hersien en sy verwantskap met Hemizygia {Benth.)

Briq. word bespreek. 'n Nuwe species, S. comptonii

Codd word beskryf en die volgende nuwe kombinasies

word gemaak; S. parviflorus var. lanceolatus ( Guerke )

Codd (=S. lanceolatus Guerke) en S. latidens ( N.E .

Br.) Codd (=Orthosiphon latidens N.E. Br.).

INDEX

Page

Hemizygia cooperi Briq 26

latidens (N.E. Br.) Ashby 26

macrophyllus ( Guerke ) Codd 27

Orthosiphon latidens N.E. Br 26

macranthus Guerke 26

Syncolostemon E. Mey. ex Benth 21

argenteus N.E. Br 3

comptonii Codd 3

concinnus N.E. Br 2

cooperi Briq 22

densiflorus Benth 25

dissitiflorus Benth 22

eriocephaius Verdoorn 25

lanceolatus Guerke 22

var. cooperi (Briq.) N.E. Br 22

var. grandiflorus N.E. Br 22

latidens {N.E. Br.) Codd 6

macranthus ( Guerke ) Ashby 6

macrophyllus Guerke 27

parviflorus E. Mey. ex Benth 2

var. dissitiflorus (Benth.) N.E. Br 22

var. lanceolatus ( Guerke ) Codd 22

var. parviflorus 22

ramulosus E. Mey. ex Benth 25

rotundifolius E. Mey. ex Benth 5

■

-

Bothalia 12,1: 29-48 (1976)

Studies in the Ericoideae. I. The genera Eremia and J.remiella

E. G. H. OLIVER*

ABSTRACT

A revision of the genus Eremia , in which seven species are recognized, and of the monotypic genus

Eremiella is presented. Both genera belong to the Ericaceae-Ericoideae and are endemic in the south-western

part of the Cape Province. The revision necessitated the inclusion of the monotypic genus Eremiopsis N.E. Br.

under Eremia. This is the first in a series on the minor genera of the Ericoideae in Southern Africa.

Resume

ETUDES SUR LES ERICOIDEAE. I. LES GENRES EREMIA ET EREMIELLA

Urie revision du genre Eremia, dans lequel sept especes sont reconnues, et du genre monotypique Eremiella

est presentee. Les deux genres appartiennent aux Ericaceae-Ericoideae et sont endemiques dans la partie sud-ouest

de la province du Cap. La revision a necessite T inclusion du genre monotypique Eremiopsis N.E. Br sous Eremia.

Cet article est le premier d'une serie sur les genres mineurs d' Ericoideae en Ajrique australe.

HISTORICAL OUTLINE

David Don proposed “An Attempt at a New

Arrangement of the Ericaceae” in 1834. In this paper

he pointed out that the generic characters in the

family are not so strongly marked, but on that account

a subdivision should not be disregarded. He dis-

approved of the large size of the genus Erica, which he

attempted to subdivide into a number of minor

groups giving each of them generic status.

Don upheld the genera Calluna and Salaxis of

Salisbury and Blaeria of Linnaeus and described the

monotypic genus Eremia basing it on Erica totta

Thunb. He separated off the rest of the species of

Erica into sixteen minor genera equivalent to the

sections used by Guthrie and Bolus in Flora Capensis

(1905). This arrangement was repeated the same year

in his brother’s work (G. Don, 1834).

Klotzsch (1838a) upheld Eremia and added E.

bartlingiana based on Drege material in Berlin.

Rach (1853), in examining the Thunberg herbarium

concurred with Klotzsch in treating E. totta and

E. bartlingiana as two distinct species. The species was

recognized by subsequent authors until N. E. Brown

(1905) rightly showed that there was no justification

for keeping it separate from E. totta. In the addenda

to his slightly earlier work, Klotzsch (1838b) described

Eremia parviflora and Eremia recurvata. The former

is now placed under Grisebachia.

The following year Bentham (1839) rearranged the

Ericaceae in his work for De Candolle’s Prodromus.

He reduced most of Don’s genera to sections of Erica

and upheld Eremia in a much enlarged form. He

placed Eremia in his subtribe Salaxidae characterized

by single ovules in each ovary cell.

Bentham divided Eremia into four sections as

follows:

Section Eremiastra characterized by a 4-celled

sessile ovary, 8 stamens and approximate bracteoles

(E. totta).

Section Poderemia characterized by a 4-celled

stipitate ovary, 8 stamens and remote bracteoles ( E .

tubercularis).

Section Micreremia characterized by a 2-celled

ovary, 8 stamens and approximate bracteoles ( E .

brevifolia, E. parviflora and E. recurvata).

Section Hexastemon characterized by a 2-celled

ovary, 6 stamens and approximate bracteoles (£.

lanata).

* Botanical Research Unit, P.O. Box 471, Stellenbosch.

This enlarged and altered treatment of the genus

brought together a heteromorphous collection of

species some of which bear no relation to each other

at specific level. It was repeated in Bentham &

Hooker (1876).

Bolus (1894) described Eremia rhodopis and placed

it in Eremia as construed by Bentham. Later Bolus

(1905) realized the close relationship between the

two species in section Poderemia and some species

in the genus Erica , section Euryloma, and proceeded

to place Eremia rhodopis and Eremia tubercularis

into Erica.

Drude (1897) recognized Eremia, but changed its

circumscription entirely by including several previously

described genera and subdividing the genus into four

sections. He retained the sections Poderemia, Eremia-

strum and Hexastemon established by Bentham and

added the section Grisebachia based on Klotzsch’s

genus. This latter section included the genera Finckea,

Acrostemon and Comocephalus described by Klotzsch.

These were subsequently removed by N. E. Brown

(1905) and placed under Grisebachia and Acrostemon

as separate genera, a view which is upheld by myself.

N. E. Brown (1905) changed Bentham’s treatment

to one without any sections and removed two of the

species included by Bentham to other genera, Eremia

lanata (Klotzsch) Benth. to Hexastemon lanatus

Klotzsch, upholding the change of Eremia tubercularis

(Salisb.) Benth. to Erica tubercularis Salisb. by Bolus

in the same work (1905). He retained Eremia parvi-

flora Klotzsch commenting that he had not seen any

material, but later (1906) included it in the synonomy

of Grisebachia eremioides MacOwan, after examining

Klotzsch’s type. This view is upheld in the present

work. He also placed Eremia bartlingiana under the

synonomy of Eremia totta.

Brown’s treatment effectively removed the dis-

cordant elements from Bentham’s Eremia thus

reducing the generic circumscription to include

species with 2- or 4-celled uniovulate ovaries and

8 stamens.

Phillips (1926) repeated the arrangement in Flora

Capensis and retained Eremia unaltered.

When dealing with specimens from the Ceres

Wildflower Show, Compton (1935) came across what

he regarded as four distinct undescribed species of

Eremia and named them E. florifera, E. peltata, E.

calycina and E. virgata. This involved no major

alterations to the generic circumscription. Compton

pointed out the close similarity between Eremia and

Erica particularly in regard to his Eremia florifera.

30

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREMIELLA

Phillips (1944) put forward his proposals for a

reclassification of the family in South Africa for the

second edition of his Genera (1951). He stressed the

character of ovule number as the principal criterion in

the subdivision of the family. He believed that the

very similar looking minor genera were more uniform

in their general appearance than the species in the

single genus Erica and should be recognized as such.

This latter statement is quite correct, but that is as

far as it goes. The variation found in Erica is almost

entirely in size, shape and colour whereas in the

minor genera the remarkable variation is in the

number of parts per floral whorl and their com-

binations.

With the above two criteria in mind, Phillips

proceeded to recircumscribe all of the genera. He

retained Eremia purely on grounds of priority and

reduced the following genera to synonomy under

Eremia: Platycalyx, Hexastemon, Arachnocalyx,

Grisebachia, Acrostemon, Simocheilus, Thoraco-

sperma and Aniserica, in fact all those genera with

uniovulate ovaries other than 1 -celled.

Phillips appeared to ignore other characters and

definite close relationships between genera. He

reduced Eremiopsis to synonomy under Scyphogyne

thus completely obscuring its close relationship with

Eremia with which it is combined in this present

paper. Further discussion of Phillip’s treatment of

the other genera will appear under the relevant

genera when published in this series.

MORPHOLOGY

In habit most of the species of Eremia are erect or

prostrate and spreading. E. totta, E. recurvata and

E. curvistyla are usually laxly spreading with E.

peltata and E. brevifolia erect and almost fastigiate.

Masson’s record of the latter at “3 to 4 feet high”

seems very unlikely. E. florifera is described by

Compton as erect. E. calycina forms a low compact

shrublet.

The branches of all species are never entirely

glabrous. There is usually some short pubescence

with another type of hairiness admixed, either long and

hispid or gland-tipped.

The leaves are all ericoid and are 3-nate in all

species except E. brevifolia in which they are 4-nate.

In most species, the leaves are erect and imbricate, but

in E. recurvata and E. curvistyla they are characteris-

tically erect-spreading to much recurved.

The flowers of all species are terminal either at the

ends of main branches or more often at the ends of

short lateral branchlets. These may then be aggregated

together to form a congested pseudospike as in

E. totta and E. curvistyla. The 3-bracteolate flowers

are usually 3-nate or as much as 10-nate in E. recur-

vata. In E. brevifolia the bracteoles are so enlarged

that the inflorescence of 4 flowers becomes involucrate

with the bracteoles giving the basic colour.

The calyx in all species is 4-lobed or 4-partite. In

E. curvistyla, E. calycina, E. florifera, E. peltata and

E. recurvata there is a distinct tubular base, which is

most marked in E. calycina. The tube is distinctly

narrow with spreading lobes in all of these except

E. recurvata. In E. totta the calyx segments are free

or very slightly joined at the base. They are entirely

free in E. brevifolia. In all species except E. brevifolia

the calyx segments are more or less equal. In this

species the type is odd in having the adaxial and

abaxial segments different in shape, but similar in

size.

The number of stamens in all specimens examined

was constantly 8 and is important in the generic

classification. In E. peltata Compton recorded

“stamens 8, sometimes but rarely 6”. This I have not

found in any of the flowers examined. The record

of Compton’s must have been a chance flower. All

stamens are free at anthesis and have slender linear

or filiform filaments. In E. recurvata the filaments are

dilated towards the base.

The anthers are all manifest or just included, the

most visible being in E. peltata. The majority of

anthers are bilobed and not bipartite with completely

free cells. Only in E. curvistyla and E. recurvata

are the anthers bipartite, but not as markedly as is

found in Eremiella or Grisebachia.

The pollen is of two types. The grains are in tetrads

in E. totta and E. florifera and single in the other

species. The former condition is typical of the

Ericaceae and is shared with Erica, the latter with

many of the other minor genera.

The ovary is either 1, 2 or 4-celled, but always

with a single subapical pendulous ovule in each cell.

In E. totta and E. florifera the ovary has 4 cells as in

Erica. In E. peltata, E. calycina, E. recurvata and

E. brevifolia the ovary always has 2 cells and is erect

and regular with an apical style. In E. curvistyla the

ovary is obliquely 1 -celled with the style arising

from the eccentric apex and curving upwards. In

one collection of this species 2-celled regular ovaries

were found in about 80% of the flowers.

As in most species of the Ericoideae, the ovary is

seated on a nectariferous disc in E. totta, E. florifera,

E. curvistyla, E. calycina and E. recurvata. This is

absent in E. peltata and E. brevifolia.

The stigma varies from simple in E. totta, E.

florifera, E. calycina and E. recurvata to slightly

capitellate in E. totta, capitate in E. brevifolia and

peltate-crateriform in E. peltata.

Mature fruits and seeds have not been seen in any

of the species. When old flowers are examined it is

found that the fruits are either very hard and nut-like

with mostly soft juicy seeds inside, if they have

developed, or they are soft and thinwalled and dis-

integrate. In the latter case soft juicy seeds occur

unlike the hard dry type occurring in Erica. An

examination of developing ovaries shows distinct

“suture” lines down the locules suggesting loculicidal

dehiscence which does not seem to occur. This

problem is met with in the majority of minor genera

and needs a considerable amount of field study to

elucidate.

DELIMITATION OF THE GENUS EREMIA

As defined in the present work the genus Eremia

is characterized by having 3 bracteoles, 4 sepals which

are free or partly fused, a 4-lobed corolla, 8 free

stamens and an ovary with 4, 2 or 1 cell with a single

ovule in each cell. The important character is the

stamen number.

When first described by D. Don in 1834, Eremia

was based on the single species E. totta with 4 unio-

vulate cells to the ovary. The uniovulate condition of

the ovary served to distinguish it from the genus

Erica and still is the only real differentiating character.

Klotzsch (1838) altered the generic circumscription

by introducing the 2-celled Eremia recurvata. Bentham

(1839) added Eremia brevifolia and proceeded to

broaden the limits by including Salisbury’s Erica

tubercularis, which has more than one ovule per cell

and also Klotszch’s Hexastemon lanatus with 6

stamens.

E. G. H. OLIVER

31

Bolus (1905) included Erica trichroma, Erica tuber-

cularis and Erica rhodopis in the section Euryloma

of Erica on the grounds that they have ovaries with

cells generally more than 1-ovuled. He stated under

Erica trichroma that the ovule number is not con-

stantly 1. He examined Niven 147 and found 12 ovules

in the 4 cells, in Masson s.n. he found 17 ovules, in

Schlechter 10278, 17 ovules and in Schlecther 7530,

6 and 7 ovules. In both Erica rhodopis and Erica

tubercularis he regarded the ovaries as 4-celled with

sometimes only one ovule per cell.

An examination of numerous flowers of my own

collection of Erica trichroma showed 12 ovules per

cell ( Oliver 4176). I also examined my collection of

Erica rhodopis ( Oliver & Palser 73) and found most of

the ovaries had 7 ovules, i.e. one cell had only 1 ovule,

and only a few had 8 ovules.

Erica trichroma and Erica tubercularis are closely

related and bear no resemblance to any of the species

of Eremia. Erica rhodopis on the other hand has a

superficial resemblance to Eremia ca/ycina, but is more

closely related to the former two species. It occurs in

the Houw Hoek to Hermanus area.

It would appear from the above that the key

character to use in separating Eremia from Erica

both of which have 4-celled ovaries is:

Ovules 4 or less per ovary Eremia

Ovules more than 4 per ovary Erica

In describing Eremia florifera Compton pointed to

its close resemblance to the section Arsace in the genus

Erica, particularly to the material that is classified

as Erica copiosa Wendl. and occurs in the Ceres

district. The ovule number in this species is, however,

at least 7 per cell as opposed to the single one in

E. florifera.

Eremia totta and E. florifera are the only 4-celled

species in the genus and with their 8 stamens are

very closely allied to the genus Erica. The overlap

between these two species and the Erica spp.

mentioned above suggests the possibility of com-

bining the two genera. This step would, however,

complicate the position regarding the relationship

between Erica and Philippia and Blaeria.

The basic ovary character of the genus Erica,

which at present contains 625 species, is 4-cells with

numerous ovules per cell. The very few exceptions to

this should in no way be used to bring about the

recircumscription of such a large genus. It will thus

be necessary in the treatment of the minor genera to

regard some of the slightly overlapping genera allied

to Erica as “genera of convenience”.

The 2-celled uniovulate erect ovary found in

Eremia peltata, E. ca/ycina, E. recurvata and E.

brevifolia is constant in all the material examined.

In nearly all the material examined belonging to

Eremiopsis curvistyla it was found that the ovary was

constantly 1 -celled, uniovulate and oblique with a

remarkable eccentric curved style. This appeared to

be a very good character for distinguishing this taxon

from all other Ericoideae. However, an examination

of Esterhuysen 29687 showed that the majority of

flowers had 2-celled uniovulate erect ovaries of the

Eremia type and a few of the Eremiopsis type. This

factor made it impossible to place the material in

either Eremia or Eremiopsis with any certainty.

The question then arose whether to keep Eremiopsis

curvistyla as a separate monotypic genus with a

partial overlap of the basic generic characters or to

incorporate it into Eremia. This had to be looked at

in the light of the generic differentiating characters

used between other genera in the subfamily. It was

decided to incorporate Eremiopsis curvistyla into

Eremia and regard it as an aberrant reduced form in

the process of evolving into a separate genus. Also,

to retain a monotypic genus there should be a very

distinct discontinuity with no overlap at all.

In Eremia a further close relationship occurs with

certain species in the genus Grisebachia and the

monotypic genus Eremiella. Grisebachia exhibits very

little variation having constantly 4-stamens and

bipartite anthers. It is a relatively natural genus of

closely related species easily distinguishable from all

other genera. There is a remarkable similarity between

Grisebachia parviflora and Eremia curvistyla and to

some extent G. minutiflora, all of which have a similar

sprawling habit and flower shape. E. curvistyla has

the bipartite anthers of the Grisebachia type, but the

8 stamens of Eremia. Undoubtedly these species had

some closer ancestral relationship in a Grisebachia-

Eremia complex.

PHYTOGEOGRAPHY

The genus Eremia is endemic in the south-western

and southern parts of the Cape Province corres-

ponding to the Western and Southern Phytogeogra-

phical Groups proposed by Weimarck (1941). There

is a conspicuous concentration of species in his

north-western centre comprising the Winterhoek and

Cedarberg Subcentres where six of the seven species

occur. E. totta, the most widespread and common

species in the genus, is spread further south as far as

French Hoek and the Hottentots-Holland in Wei-

marck’s South-western Centre. Of particular interest

is the northern record of this species from the

Heerenlogementberg where very few Ericaceae have

been recorded. The species also occurs in the “island

floras” on the Piquetberg mountains, Riebeek

Kasteel and Paarl Mountain and on the Paardeberg

near Malmesbury. Of the other species in this area,

E. peltata is of interest in extending out of the

Bokkeveld onto the Bonteberg near Touws River

where isolated patches of Cape flora occur on the

mountains of the eastern end of the Little Karoo.

The only representative of the genus outside the

above area is E. brevifolia which is restricted to a

single locality, Attaquaskloof, in the Outeniqua

Mountains about 200 km east of the rest of the genus.

This disjunction should be looked at in conjunction

with the occurrence slightly further east of the closely

related monotypic genus Eremiella.

All of the species occur on dry stony soils of the

Table Mountain Series in the Cape System associated

with other members of typical mountain fynbos.

Most of them grow on dry rocky slopes, but E.

ca/ycina is found on dry sandy flats of the Cold

Bokkeveld north of Ceres.

EREMIA

Eremia D. Don emend. E. G. H. Oliver. Eremia

D. Don in Edin. New. Phil. Journ. 17: 156 (1834);

G. Don, Gen. Syst. 3: 828 (1834); Klotzsch in

Linnaea 12: 218 (1838); Benth. in DC., Prodr. 7: 699

(1839) pro parte; et in Benth. & Hook, f., Gen. PI.

2: 592 (1876); Drude in Pflanzenfam. 4,1: 63 (1897)

pro parte; N.E. Br. in FI. Cap. 4,1: 332 (1905);

Phill., Gen. ed. 1, 460 (1926); Compton in J.S. Afr.

Bot. 1: 150 (1935); Phill. in J.S. Afr. Bot. 10: 71

(1944) pro parte minore; et Gen. ed. 2, 560 (1951).

Type species: E. totta (Thunb.) D. Don.

Eremiopsis N.E. Br. in FI. Cap. 4,1: 390 (1906).

3:

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREM1ELLA

The generic name is derived from the Greek word,

eremos, meaning single or solitary and refers to the

solitary ovule in each cell of the ovary.

Perennial woody shrublets, erect up to 30 cm or

prostrate and spreading. Leaves 3-nate, in one

species 4-nate, erect, imbricate to spreading and

recurved. Flowers terminal usually on short lateral

branchlets, often forming congested pseudo-spikes.

Bracteoles 3, mostly approximate, small and incon-

spicuous to large forming an involucre around the

inflorescence, glabrous rarely pubescent, ciliate. Calyx

4-partite or lobed, small to enlarged and conspicuous;

segments mostly equal, sometimes in two slightly

dissimilar ranks, in one species slightly unequal,

glabrous rarely pubescent, ciliate with simple, gland-

tipped or plumose hairs, in one species sparsely villous

never lanate. Corolla 4-lobed, urceolate, campanulate

or cyathiform, glabrous rarely puberulous. Stamens

8, free, included or manifest. Anthers bipartite or

bilobed, aristate or muticous. Pollen in tetrads or

single-grained. Ovary 2- or 4-celled with a single

subapical pendulous ovule in each cell, ovoid, or

obliquely 1 -celled with a single oblique subapical

ovule. Nectariferous disc present or absent. Style

exserted or included, apical and straight erect in 2-

or 4-celled ovaries or eccentric and crooked in 1 -celled

oblique ovaries. Stigma simple, capitate or peltate-

crateriform. Fruit a hard apparently indehiscent nut

or soft and dry, breaking by decay.

The genus is divided into 2 subgenera on the ovary

characters.

KEY TO THE SPECIES

Ovary 4-celled Subgenus: Eremia

Corolla 4-Iobed and swollen in lower half; sepals free, 2-3 mm long; anthers muticous. A. E. totta

Corolla rounded and narrow in lower half; sepals united at base, up to 0,9 mm long; anthers

aristate 2. E. florifera

Ovary 2- or 1 -celled Subgenus: Metagyne

Ovary 2-celIed; style apical:

Stigma peltate-crateriform 3. E.peltata

Stigma simple or capitate:

Anthers aristate, scabrid, not bearded:

Leaves more or less erect, adpressed 4. E. calycina

Leaves spreading-recurved:

Calyx lobes orbicular to broadly elliptic, cilia much shorter than the width of the

lobe 5. E. curvistyla

Calyx lobes narrowly ovate to ovate-acute, cilia longer than the width of the

lobe 6. E. recurvata

Anthers muticous, smooth, bearded in front 7. E. brevifolia

Ovary l-celled; style eccentrically placed 5. E. curvistyla

E. G. H. OLIVER

33

Subgenus Eremia

Ovary 4-celled with a single ovule in each cell.

Pollen in tetrads.

Type: E. totta (Thunb.) D.Don.

The subgenus contains two species, E. totta and

E. florifera, which possess the characters used in the

original circumscription of the genus.

1. Eremia totta (Thunb.) D.Don in Edin. New

Phil. Journ. 17: 156 (1834); Klotzsch in Linnaea

12: 218 (1838) pro parte; Benth. in D.C., Prodr.

7: 699 (1839) pro parte; N.E. Br. in FI. Cap, 4, 1 : 332

(1905). Type: Cape. Thunberg (a) no. 9437 (UPS).

Erica totta Thunb., Diss. Bot. (Erica) 18 (1785);

Prodr. 70 (1794); Diss. Acad. (Erica) 2: 215 (1800).

FI. Cap. 348 (1811); Bartl. in Linnaea 7: 647 (1832)

pro parte.

Euremia totta (Thunb.) Rach. in Linnaea 26: 789

(1855).

Erica ferox Salisb. in Trans. Linn. Soc. 6: 324

(1802), nom. illegit. Type as for E. totta.

Erica pectinata Bartl. in Linnaea 7: 647 (1832).

Type: Tulbagh Kloof, Ludwig & Beil s.n. (B, holo.|;

isos?).

Eremia bartlingiana Klotzsch in Linnaea 12: 218

(1838); Benth. in DC., Prodr.: 699 (1839). Type: Du

Toits Kloof, Drege s.n. (B, holo.f; K!; P!).

Euremia bartlingiana (Klotzsch) Rach in Linnaea

26: 789 (1855).

Eremia totta var. bartlingiana (Klotzsch) N.E.Br. in

FI. Cap. 4,1: 333 (1905).

Low, spreading shrublet to 30 cm high, rarely

higher. Branches stout, minutely pubescent all over

sometimes sparsely so, with sparse or dense, long,

spreading hairs which are minutely hispid mostly

more so towards their bases and with few to many

simple shorter gland-tipped hairs admixed. Leaves

3-nate, spreading or reflexed but curved upwards,

2-3 mm long without the petiole, angular when

dried, narrowly elliptic to linear-elliptic, minutely

pubescent all over, rarely glabrous, with long, white,

spreading hairs becoming subechinate on the sides

and abaxial surface, hairs often hispid when young;

petiole usually adpressed, up to 0,5 mm long,

pubescent or glabrous, with short hairs on the margins.

Flowers terminal, (2)3(4)-nate on ends of minute

axillary branchlets clustered together at the ends of

short lateral branchlets to give a congested pseudo-

spike; pedicels 0,5-1, 3 mm long, glabrous or

puberulous, sometimes shortly glandular pubescent;

bracteoles 3, adpressed to the calyx 0,7-2, 1 mm long

and up to 2 mm broad, elliptic to ovate to depressed

ovate, acute, rarely obtuse, keel-tipped, glabrous or

occasionally puberulous, serrulate-ciliate or fimbriate

with mostly hispid cilia, not glandular, occasionally

with a few stout hairs on the keel-tip, white. Calyx

equally 4-partite or occasionally slightly joined at the

base, reaching from halfway to completely up the

corolla-tube; segments 1,7-3, 5 mm long and 1,4-

2 mm broad, elliptic-oblong to broadly elliptic,

sometimes obovate, mostly obtuse and cucullate, with

or rarely without a keel-tip, glabrous or puberulous,

serrulate-ciliate or fimbrate, the “cilia” often minutely

hispid and gland-tipped mostly towards the apex

where the glands become subsessile, occasionally

with a few hairs on the keel-tip. Corolla 4-lobed,

2, 5-3, 9 mm long and up to 2,6 mm wide, mostly

urceolate with varying degrees of narrow neck from

a large inflated markedly 4-winged base, the wings

alternating with the sepals, rarely constricted below

the erect broad rounded lobes, glabrous or rarely

minutely puberulous, white. Stamens 8, free, included;

filaments filiform, glabrous, up to 1,8 mm long;

anthers 0,9x0, 2 mm oblong, bilobed to bipartite

dorsifixed near the base, smooth or slightly scabrid,

muticous; pore about half length of the cell; pollen

in tetrads. Ovary 4-celled, with a single pendulous

subapical ovule in each cell, 0, 8x1,1 mm, broadly

and obtusely conical to ellipsoid 4-angled, glabrous

to densely lanate mainly at the apex; style up to 3 mm

long; included or shortly exserted, filiform, glabrous,

stigma simple to subcapitellate. FiG. 2.

A low, spreading shrublet, rarely erect, occurring

frequently on dry mountain slopes from the

Vanrhynsdorp district south-wards to the Caledon

district, the most widespread and commonest species

in the genus.

Cape. — 3118 (Vanrhynsdorp): Heerenlogementberg, 300-

600 m (-DC), Sept. 1962, Taylor 3946 (STE). 3218 (Clanwilliam):

Pilaarsberg, Pakhuis, 910 m (-BB), Sept. 1967, Kerfoot 5918

(NBG); Schimmelberg (-BD), Oct. 1939, Pillans 9114 (BOL);

Redlinghuys, 180 m (-CB), Oct. 1958, Acocks 19794 (K; PRE;

Z); Gruyskop, Piquetberg (-DA), Nov. 1934, Pillans 7214

(BOL; K); Kapteins Mountain, 1 036 m (-DA), Oct. 1935,

Pillans 7772 (BOL); Kapteinskloof (-DA/DC), Sept. 1955,

Van Niekerk 625 (BOL; STE); Grey's Pass (-DB), Oct. 1925,

Levyns 1361 (CT); Pikenier's Pass (-DB), Nov. 1910, Pillans

5124 (BOL; K); Zebrakop, 1 450 m (-DB), Nov. 1934 Pillans

7567 (BOL); Waboom, Piquetberg, 760 m (-DB), Oct. 1963,

Taylor 5352 (K; STE); Mouton Valley, Piquetberg, 760 m

(-DC), Oct. 1922, Marlotlt 11504 (PRE; STE); Versveld Pass

(-DD), Nov. 1934, Pillans 7166 (BOL); Piquetberg between

Versveld Pass and Zebrakop, 762 m (-DD), Sept. 1962, Taylor

3902 (STE); Piquetberg Mountain, 600-900 m (-DC/E)D),

Nov. 1828, Drege s.n. (P); 578 m, Oct. 1892, Guthrie 2655

(BOL); 760 m, Nov. 1951, Johnson 291 (NBG; STE); Sept.

1927, Levyns 2167 (CT); Sept. 1949, Martin 246 (NBG); Nov.

1951, Martin 906 (NBG). 3219 (Wuppertal): Krakadouw 914 m

(-AA), Oct. 1897, Bodkin s.n. (BOL); Pakhuis Pass (-AA),

Sept. 1940, Bond 588 (NBG); Koudeberg near Wuppertal,

1 100 m (-AA), Aug. 1896, Schlechter 8750 (BM; K; PRE);

Pakhuis Pass at Leipoldt’s Grave (-AA), Nov. 1971, Schlieben &

Ellis 12453 (PRE; STE); Charity Peak, 1 066 m (-AA), Sept.

1936, Thorne in SAM 54845 (SAM); Niewoudt Pass, 610 m

(-AC), Sept. 1934, Compton 4920 (BOL; NBG); Uitkyk Peak,

1 520 m (-AC), Dec. 1941, Esterhuysen 7369 (BOL); Algeria

(-AC), Oct. 1930, Galpin 10562 (PRE); Scorpion’s Poort,

1 220 m (-AC), Oct. 1923, Pocock 497 (PRE, STE); Sneeuberg

hut area (-AC), Dec. 1964, Taylor 6175 (STE); Uitkyk Pass

(-AC), Sept. 1937, Wall 42 (S); Blaawberg (-AA/AC), Dec.

1830, Drege s.n. (P); Middelberg Pass, 610 m (-CA), Nov.

1955, Baker 828 (BM); Elandskloof (-CA), Sept. 1944, Compton

16129 (NBG; STE); 1 066 m. Sept. 1936, Levyns 5770 (CT);

Warmbaths, Citrusdal (-CA), Sept. 1911, Stephens 6891 (K;

NH; PRE); Stephens 7038 (BM; K; SAM; STE); Skoongesig,

Bokkeveld (-CC), Aug. 1969, Hanekom 1270 (K; PRE);

Cardouw Pass, 910 m (-CC), Nov. 1951, Maguire 1208 (NBG;

STE); Onderboskloof (-CC), Sept. 1958, Middlemost 1971

(NBG; STE); Berghof, Porterville mountains, 790 m (-CC),

Sept. 1972, Oliver 3937 (MO; PRE; STE). Without precise

locality: Clanwilliam district, Oct. -Feb., Leipoldt 212 (BOL;

SAM); Cedarberg, Aug. 1896, Mann sub Marloth 11377

(PRE); Cedarberg, Oct. 1923, Pocock 116 (STE). 3318 (Cape

Town); Riebeek Kasteel (-BD), Oct. 1968, Bavliss 4330 (MO;

NBG); 600 m, Oct. 1968, Marsh 1032 (K; PRE; STE); Niven

69 (G-DC); Niven 83 (K); Sept. 1929, Pillans 7009 (BOL);

Paardeberg (-DB), Oct., Beil in SAM 41319 (SAM); Paarl

Mountain (-DB), Bolus 2948 (K); 610 m, Oct. 1942, Henderson

1190 (NBG); at Gordon Rock, Sept. 1961, Jordaan 1288 (STE);

Aug. 1883, Wilms 3403 (K); Swartboskloof (-DD), Sept. 1936,

Borcherds 418 (STE); Aug. 1961, Van der Merwe 26-71 (PRE);

Stellenbosch Mountain (-DD), Sept. 1917, Garside 1009 (K);

Sept. 1946, Rehm s.n. (M); June 1946, Strey 448 (PRE);

Stellenbosch (-DD), Oct. 1891, Guthrie 2372 (BOL); Harvey s.n.

(E); Jonkershoek (-DD), Sept. 1928 Markotter 2018 (STE);

Oct. 1846, Prior s.n. (K; PRE); Elsenberg, Warwick Farm,

(-DD), Oct. 1938, Penfold 154 (NBG). 3319 (Worcester);

Winterhoek, 700m (-AA), Nov. 1879, Bolus 5192 (BM; BOL;

Z); Inkruip, Witsenberg (-AA), Oct. 1945, Esterhuysen 23449

(BOL; STE); Saronsberg, 450-610 m (-AA), May 1956,

Esterhuysen 25812 (BOL; STE); Little Winterhoek, 1 200 m

(-AA), Oct. 1884, Marloth 500 (PRE); Sneeugat (-AA),

Nov. 1916, Phillips 1816 (SAM); Oct. 1934, Thorne in SAM

51252 (SAM; STE); near Saron, 550 m (-AA), Oct. 1896,

Schlechter 10687 ( BM; E; G; K; MO; P; S; W; Z); Ertjiesland-

kloof (-AB), Sept. 1944, Compton 16099 (NBG); Gydo Pass

34

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREM1ELLA

Fig. 2. — Eremia totta. 1, flower

xlO. 2, corolla, xlO. 3,

bracetoles: a, median; b,

laterals, all x20. 4, sepal,

x20. 5, anther, side, front

and back views, all x20. 6,

androecium and gynoecium*

x 10. All drawn from the

type, Thunberg 9437 (UPS).

7, median bracteole, x20,

drawn from Taylor 6175

(STE). 8, sepal, X 20, drawn

from Hutchinson 614 (PRE).

9, gynoecium, xlO, drawn

from Oliver 4064 (STE). 10,

hairs found on an average

young branch, X 20.

(-AB), Oct. 1928, Hutchinson 1009 (BOL; K); Witsenberg

Pass (-AC), Dec. 1919, Andreae 129 (STE); Witsenberg (-AC),

Oct. 1927, Barnard in SAM 20961 (SAM); Witsenberg (-AC),

Dec., Zeyher 1116 (BM; G; K; P; SAM; STE); Oct., Zeyher

in SAM 41316 (SAM); Tulbagh Waterfall, 450 m (-AC),

Oct. 1882, Bolus 5303 (BOL); 300-600 m, Nov., Ecklon &

Zeyher s.n. (K); Oct. 1928, Herre in STE 8976 (STE); Sept.

1946, Rehm s.n. (M); Sept. 1892, Schlechter 1439 (BM; K);

300 m, Sept. 1903, Thode in STE 5465 (STE); Ceres Peak

(-AD), Oct. 1933, Acock s 1829 & 1838 (S); Gorge W. of Ceres

(-AD), Sept. 1928 Hutchinson 614 (BM; BOL; K; PRE);

Ceres (AD), Oct. 1924, Levyns 1078 (CT); Hill close to Ceres,

500 m (-AD), Aug. 1921, Marloth 10353 (PRE; STE); Mosterts-

hoek, 1 600 m (-AD), Oct. 1894, Marloth 2002 (PRE); Schurfte-

berg, Ceres (-AD), Nov. 1941, Pi/Ians 9696 (BOL); Witsenberg

Vlakte (-A?), Oct. 1941, Walgate 392 (NBG); Matroosberg,

(-BC), Oct. 1893, Marloth 2014 (STE); Hex River Valley (-BC),

Sept. 1881, Tyson 700 (BOL; K); Bainskloof (-CA), May

1939, Bond s.n. (NBG); Oct., Ecklon in SAM 41320 (SAM);

Oct. 1930, Fries, Norlindh & Weimarck 1713 (S); Sept. 1938,

Hafstrom & Acocks 1053 (PRE; S); Sept. 1936, Lindeberg s.n.

(S); 550 m, Sept. 1885, Marloth 647 (PRE); Sept. 1931,

Neethling s.n. (STE); near Stoneman Station, Bainskloof (-CA),

July 1956, Rycroft 1947 (NBG); Seven Sisters Mountain Paarl

(-CA), Oct. 1943, Esterhuysen 9027 (BOL); Molenaar’s Peak

(-CA), Oct. 1947, Stokoe in SAM 62525 (SAM); Dutoitskloof,

300-450 m (CA), Sept. 1828, Drege s.n. (K; P); Dutoitskloof

Tunnel ( -CA), Oct. 1951, Maguire 1141 (NBG); Peak north of

Chavonnesberg, 1 220 m (-CB), Dec. 1921, Andreae S2<?(STE);

Chavonnesberg (-CB), 1922, Stokoe in PRE 32029 (PRE);

Veld Reserve, Worcester, (-CB), July 1934, Van Breda 33

(PRE); Franschhoek (-CC), Oct. 1946, Barker 4159 (NBG);

550 m. Sept. 1895, Bolus s.n. (PRE); Dec. 1934, Hafstrom s.n.

(LD; S); 300 m, April 1887, MacOwan sub Herb. Norm. 756

(BM; BOL; G; K; P; SAM; UPS; W; Z); MacOwan 2826

(E; SAM); Dec. 1933, Meebold 14960 (M); Oct. 1846, Prior s.n.

(K); Franschhoek Pass, 914 m (-CC), Sept. 1935, Compton

5673 (BOL); April 1931, Levyns 3421 (CT); Franschhoek, foot

of mountains behind Kriel (-CC), Oct. 1913, Phillips 1233

(SAM); Berg River Hoek (-CC), Sept. 1946, Compton 18317

(NBG); Sept. 1946, Leighton 2053 (PRE); Zachariashoek,

426 m (-CC), Sept. 1968, Kruger 704 (STE); 730 m, Oct. 1972,

Oliver 4064 (K; MO; PRE; STE); 1 220 m, Aug. 1961, Van

der Merwe 818 (PRE); Wemmershoek Mountains, N. of

Paardekop, 1 220 m (-CC), Oct. 1943, Wasserfall 492 (NBG);

Wemmershoek Mountains, Turkkloof, 457 m (-CC), Oct. 1943

Wasserfall 508 (NBG); Scherpenheuwel (-DA), Sept. 1951,

Barker 7522 (NBG; STE); Wildepaardeberg (-DC), Stokoe,

6255 (BOL). Without precise locality: Worcester, 1859, Cooper

1703 (BM; K; W; Z); Drakenstein et Dutoitskloof, 760. 1 060 m,

D'-ege s.n. (G-DC); Worcester, Tulbaghkloof, Tulbagh, foot of

Winterhoek, Witsenberg and at Vogelvlei, Sept., Ecklon and

Zeyher s.n. (E; K; LD; MO; P; S; UPS; Z); Tulbagh, Oct.

1890, Guthrie 2078 (BOL), Tulbagh, Oct. 1901, Kassner 1127

(E); Tulbagh, Pappe s.n. (K); Ceres Nov. 1933 Meebold 14961

(M); Worcester mountains, Oct., Zeyher s.n. (SAM). 3418

(Simonstown): Helderberg, 610 m (-BB), Sept. 1907, Diimmer

565 (E); Sir Lowry’s Pass (-BB), Sept. 1936, Hafstrom and

Lindeberg s.n. (S); Lourensford, 250 m (-BB), Aug. 1948.

Parker 4338, (K; NBG). 3419 (Caledon): Flats E, of Viljoen’s

Pass (-AA), Sept. 1949, Davis in SAM 62547 (BM; SAM);

Stokoe in SAM 62546 (SAM); Boschmanskloof, Elgin area,

(-AA?), Oct. 1948, Stokoe s.n. (SAM; STE). Without local-

ity: Auge s.n. (BM); Drege s.n. (BM; G; LD; MO; P; S; W);

Ecklon s.n. (M; W); Admiral Grey s.n. (K; S); Gueinzius 262

(G; W); Gueinzius s.n. (S); Harvey s.n. (K); Masson s.n. (BM);

Niven s.n. (K); Roxburgh s.n. (G; K); Herb. Salisb. s.n. (K);

Scholl 271 (W); Thunberg a (3 (UPS).

E. G. H. OLIVER

35

Fig. 3. — Distribution of Eremia

totta. The variation in the

flowers for a selection of

specimens is shown. 1, Jor-

daan 1288. 2, Schlechter

10687. 3, Pillans 7214 4,

Acocks 19794, 5, Taylor

3946. 6, Taylor 5352. 7,

Schlieben & Ellis 12453.

8, Taylor 6175. 9, Hanekom

1270. 10, Oliver 3937. 11,

Barker 1522. 12, Oliver 4064.

13, Prior s.n. (Jonkershoek).

Allx2.

Fig. 4. — Eremia florifera. Twig, xl,5. 1, flower, Xl8. 2, corolla, Xl8. 3, median bracteole, Xl8.

4, calyx, ventral view, Xl8. 5, stamen, side view, Xl8. 6, anther, back view, x36. 7, anther,

side view, x36. 8, gynoecium, Xl8. 9, gynoecium, cut longitudinally, xl8. 10, ovary, cut

transversely, x36. 1 1, whorl of leaves, x 18. All drawn from the holotype, Compton 4927 (BOL).

36

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREMIELLA

E. totta is distinct in the genus for its large white

flowers which turn yellow-brown when dried. Florally

it is easily distinguishable in the subgenus by its large

free sepals and bracteoles.

Originally described as a species of Erica by

Thunberg the species formed the type of Don’s new

genus Eremia. Despite this latter fact E. totta is not

typical of the species included under Eremia in the

present revision.

Several authors found grounds for splitting off

additional species from E. totta. Bartling described

Erica pectinata and Klotzsch Eremia bartlingiana.

These were both based on variations in corolla shape

and hairiness of the ovary, characters which have

subsequently been found to be variable and of no

taxonomic significance. N. E. Brown reduced the

latter species to a variety of E. totta, but 1 have not

upheld this.

There is much variation in the hairs on the ovary

from very crisped lanate to glabrous, sometimes

with only one or two long hairs present. In his type

description, Thunberg makes no mention of the

ovary. There are two sheets in his herbarium, a and /?,

and N. E. Brown states that both specimens on

sheet a have lanate ovaries, while specimen /? has

only a few hairs. I have chosen the first as the lectotype.

The variation in the pubescence on the sepals and

bracteoles appears to be randomly distributed and

not correlated with other characters. This is also true

of the sepals and bracteoles which vary in shape and

size and in their relationship to the length of the

corolla. In Schlieben and Ellis 12453 the sepals are

broadly ovate and less than half the length of the

corolla-tube whereas in Schlechter 10687 they are

elliptic and as long as the corolla-tube (Fig. 3).

E. totta is the most widespread and common

species in the genus, occurring in dry sandy or

rocky areas from the Cedarberg to the Stellenbosch

mountains. It is one of the few Ericaceae which have

been recorded from the “island” floras of Paarl

Mountain, Riebeek’s Kasteel and Paardeberg. A very

unusual and interesting record is Taylor 3946 from

the Heerenlogementberg in the Vredendal district.

2. Eremia florifera Compton in J.S. Afr. Bot.

1: 149 (1935). Type: Ceres Wildflower Show, Oct.

1934, Compton 4927 (BOL!).

Erect, much-branched shrublet. Branches slender,

puberulous, internodes more or less elongate;

branchlets lateral, fasciculate. Leaves 3-nate, erect-

spreading, linear-oblong, up to 3,0 mm long with

petiole 0,6 mm long, obtuse, sulcate, setose when

young, glabrescent. Flowers numerous, in small

terminal clusters or clumped laterally on short

branchlets; pedicels up to 0,7 mm long, glabrous;

bracteoles more or less median, subequal to unequal,

the middle up to 0,9 mm long the laterals 0,6 mm,

all oblong, scarious, glabrous, shortly ciliate, some-

times keel-tipped. Calyx 4-lobed to about two-thirds

the way down, glabrous; tube 0,3 mm long; lobes

ovate almost angular-ovate, slightly spreading,

0,6 mm long, 0,3 mm broad, shortly ciliate. Corolla

1,7 mm long, cyathiform, slightly contracted at the

base, glabrous, pink; tube 1 ,3 mm long; lobes 0,4 mm

long, broad, obtuse, entire. Stamens 8, free, included;

filaments filiform, 0,8 mm long; anthers 0,4 mm long,

ovoid, smooth to minutely scabrid, dark brown, with

long pale scabrid awns; pore about third to half the

length of the cell; pollen in tetrads. Ovary 4-celled,

with a single subapical pendulous ovule per cell,

oblong-ovoid 0,4 mm long, puberulous; style short

thick 0,7 mm long; stigma included, simple obconic.

F;G. 4, reproduced from Compton’s original drawing.

Cape.— Without precise locality; Ceres Wildflower Show,

Oct. 1934, Compton 4927 (BOL).

E. florifera is placed with E. totta in the subgenus

Eremia on account of the possession of a 4-celled

ovary and pollen grains in tetrads. It can easily be

distinguished by its much smaller flowers, small

inconspicuous bracteoles and sepals, cyathiform

corolla and aristate anthers.

Compton (1935) drew attention to the close

resemblance of this species to members of the section

Arsace in the genus Erica, especially to what Guthrie

and Bolus regard as Erica copiosa Wendl. var.

longicauda H.Bol. The only basic difference between

E. florifera and the Erica spp. is in the number of

ovules. In the latter the number of ovules is 7-8

per cell.

The relationship between E. florifera and the genus

Erica is in fact closer than its relationship with

E. totta. It would seem highly probable that these

two species evolved from ancestral Erica stock

completely independently.

No locality can be given for the species as it has

only been recorded once at the Ceres Wildflower

Show in October, 1934. At the flower shows organized

in some centres of the south-western Cape, the

majority of exhibits are collected in the local district,

but there is no guarantee of this as a record. The

Ceres District, however, fits into the distribution of

the genus with 5 species occurring there.

Subgenus Metagyne E. G. H. Oliver subgen. nov.

ovario bicellulare vel unicellulare, ovulo uno in

quoque cellulo, pollinis granis singulis.

Ovary 2-celled or 1 -celled with a single ovule in

each cell, pollen grains single.

Type: E. recurvata Klotzsch.

The subgenus contains five species: E. peltata,

E. calycina, E. recurvata and E. brevifolia all with

erect, 2-celled ovaries and E. curvistyla with 1 -celled,

oblique ovaries, rarely 2-celled and erect.

3. Eremia peltata Compton in J.S. Afr. Bot. 1 : 147

(1935). Type: Ceres Wildflower Show, Compton

4921 (BOL!).

Erect, much-branched shrublet about 30 cm high.

Branches slender, 3-angled when young, grey-

pubescent with spreading to deflexed hairs, becoming

glabrous. Leaves 3-nate, up to 3 mm long and 0,8 mm

wide including the very short petiole, erect, narrowly

elliptic to occasionally ovate, obtuse, sulcate,

glabrous, slightly puberulous on adaxial surface,

minutely ciliate and with sessile glands admixed.

Flowers 1- to 6-nate, mostly 3-nate, terminal, erect or

slightly nodding; pedicel 1,2 mm long, glabrous;

bracteoles 3, scattered on pedicel but laterals usually

median, the middle one remote, up to 1,2 mm long,

ovate or laterals narrowly elliptic and often assymetric,

all carinate, glabrous, scarious, shortly ciliate with

sessile glands admixed. Calyx 4-lobed divided to

two-thirds the way down, campanulate, scarious,

glabrous, pink; tube up to 0,5 mm long, subquadrate;

lobes up to 1,2 mm long and broad, ovate to very

broadly ovate or elliptic, obtuse, subcucullate, keel-

tipped, shortly ciliate and with sessile glands admixed.

Corolla 4-lobed, up to 2,2 mm long, glabrous, pink;

tube up to 1,5 mm long, tubular to narrowly obconic

at the base then broadly obconic; lobes up to 0,7 mm

long and 1,5 mm broad, broadly orbicular,

obtuse, erect or slightly spreading, minutely crenulate.

Stamens 8, very rarely 6, free; filaments straight,

E. G. H. OLIVER

37

Fig. 5. — Eremia peltata. Twig, xl,6. 1, flower. 2, flower with bracteoles and calyx removed.

3, median bracteole. 4, calyx, ventral view. 5, sepal, inner surface. 6, stamen, back view. 7,

stamen, side view. 8, gynoecium. 9, gynoecium, cut longitudinally. 10, whorl of leaves. All drawn

Xl8 from the holotype, Compton 4921 (BOL).

linear, 1 ,5 mm long, glabrous; anthers up to 1,2 mm

long, oblong-ellipsoid, sometimes with a prognathous

base and colourless hairs, partly exserted, bipartite,

dorsifixed near the base, scabrid, dark brown

muticous; pollen grains single. Ovary 2-celled with a

single pendulous subapical ovule in each cell, up to

0,7 mm long, ovoid, compressed, puberulous on

upper half with or without a very small nectariferous

disc; style up to 3 mm long, slender, glabrous; stigma

far exserted, peltate-crateriform with slight lobes

inside, red. Fig. 5, reproduced from Compton’s

original drawing.

A small erect shrublet in habit, the species occurs on

dry sandy slopes in the southern part of the Ceres

district.

Cape. — 3319 (Worcester): South Cold Bokkeveld mountains

near Sandberg (-AB), Oct. 1940, Bond 653 (BOL; K; NBG;

PRE); Esterhuysen 3461 (BOL); Gydo, 1 220 m (-AB), Nov.

1946, Compton 18748 (NBG); 13 km beyond Gydo Pass (-AB)

Oct. 1928, Hutchinson 1044 (BOL; K); Eselfontein (-AD), Oct.

1953, Esterhuysen 21876 (BOL); 610 m, Aug. 1958,

Esterhuysen 27858 (BOL); Baviaansberg, 1 220 m (-BA), Nov.

1962, Esterhuysen 29848 (BOL); Jan. 1956, Stokoe in SAM

68392 (SAM); N. aspect on Matroosberg, 1 220 m (-BC), Sept.

1924, Levyns 957 (BOL; CT); Welvaart at base of Sanddrift

Peaks, 990 m (-BC), Oct. 1974, Oliver 5086 (B; BM; BOL; C;

E; G; K; M; MO; NBG; P; PRE; S; STE; W; Z); Conical

Peak (-BC), Dec. 1940, Stokoe 7716 (BOL); Bonteberg, Eiken-

boschhoek, 1 220 m (-BD), Nov. 1940, Compton 9920 (NBG);

Esterhuysen 3778 (BOL). Without precise locality: Ceres

Wildflower Show, Oct. 1925, Pillans in BOL 18421 (BOL);

Oct. 1934, Compton 4921 (BOL).

E. peltata is a very distinct species easily

recognizable in the genus by its peltate-crateriform

stigma.

In some characters it is similar to E. brevifolia.

Both species are odd in not having the typical ericoid

nectariferous disc below the ovary. This, coupled

with the dull greenish white colour of their flowers,

manifest to semi-exserted anthers with large pores

and their stigmas, peltate in E. peltata and capitate in

E. brevifolia, suggests that they are both wind

pollinated. All other species in the genus are either

distinctly white or pink with simple stigmas.

The species is fairly uniform over its distribution

range. The only noticeable variation occurs in the

anthers, which vary in shape sometimes in the same

flower. They sometimes have a remarkable prog-

nathous base and occasional long colourless hairs of

the type found in E. brevifolia. Compton recorded

that there were 8 stamens “sometimes but rarely 6”.

This I have not seen in any of the specimens I have

examined.

38

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREMIELLA

E. peltata occurs in a distinct part of the distribution

range of the genus in the southern and eastern parts

of the Ceres district, where it grows on dry sandy,

stony mountain slopes. Only in the Cold Bokkeveld

around Gydo and Sandberg does it overlap with

other species. The species occurs on the Bonteberg

north of Touws River where islands of the Cape

Flora occur in karroid vegetation types.

The species flowers from September to December.

The exact locality of the type specimen is unfor-

tunately not known as it came from the Ceres Wild-

flower Show.

4. Eremia calycina Compton in J.S. Afr. Bot.

1: 146 (1935). Type: Cold Bokkeveld, Rosendal,

Compton 4939 (BOL!).

E. virgata Compton in J.S. Afr. Bot. 1: 148 (1935). Type:

Ceres Wildflower Show, Compton 4933 (BOL!).

Erect, compact or diffuse shrublet up to 30 cm.

Branches slender, virgate or flexuose, grey-puberulous

with crisped hairs when young. Leaves 3-nate, 1,5-

2 mm long and 0,7 mm broad, imbricate or shorter

than the internodes, erect, adpressed, oblong-elliptic,

slightly hooded at apex and gibbous at the base,

sulcate, glabrous, ciliate and with sessile glands when

young. Flowers 1- to 5-nate, terminal on main

branches or short lateral branchlets, deflexed or

subcernuous; pedicels up to 0,5 mm long; bracteoles

3, subbasal, subequal, up to 0,8 mm long, erect,

oblong, obtuse, glabrous, ciliate mainly towards

apex, pink. Calyx 4-lobed, campanulate, scarious,

glabrous, up to 1,9 mm long, divided from half to

two-thirds; tube up to 1 mm long and 1 mm diameter;

lobes up to 1 mm long, the outer two up to 1,7 mm

broad, the inner two to 1,5 mm, broadly orbiculate,

indexed in upper half, broadly hooded, keeled near

top, rigidly scarious, pink, margins crenulate to

serrulate, sometimes ciliate and with a few sessile

glands admixed. Corolla 4-lobed, up to 2,7 mm long,

obconic to obconic-campanulate, glabrous, pink; tube

up to 1,7 mm long; lobes very broad, up to 1 mm

long and 1,2 mm broad, obtuse, crenulate, erect.

Stamens 8 free; filaments filiform, glabrous, bent at

apex, up to 1,6 mm long; anthers manifest to sub-

included, up to 0,5 mm long, ovoid, bipartite,

scabrid, attached dorsally near the base, aristate to

subcristate; pore half length of cell; appendages up to

0,5 mm long, pointing downwards, free or slightly

decurrent; pollen grains single. Ovary 2-celled, with

single pendulous subapical ovule in each cell, up to

0,9 mm long and 0,6 mm broad, ovoid, compressed,

subacute, puberulous at top, with a nectariferous

disc; style filiform, up to 2,5 mm long; stigma far

exserted, simple sometimes slightly lobed. Fig. 7,

reproduced from Compton’s drawing (Compton l.c.

for E. virgata).

A low-growing shrublet compact or diffuse in habit

occurring on dry open sandy flats in the Cold Bokke-

veld region of the Ceres district.

Cape. — 3219 (Wuppertal). Elandskloof mountains (-CA),

Sept. 1936, Compton 6661 (BOL; NBG; PRE); Elandskloof,

sandflats, 910 m (-CA) Sept. 1944, Compton 16131 (NBG);

Sept. 1936, Lewis in BOL 21999 (BOL); De Straat, 790 m

(-CA), Oct. 1972, Oliver 4025 (K; PRE; STE; W); Oliver 4026

(PRE; STE); Wagenboomsrivier, 910 m (-CA), Sept. 1936,

Compton 6663 (NBG); 838 m, Oct. 1972, Oliver 4010 (B; BM;

E; G; K; MEL; MO; NBG; P; PRE; S; STE); Rosendal,

910 m (-CD), Sept. 1934, Compton 4939 (BOL). 3319 (Wor-

cester) Gydo, 1 066 m (-AB), Nov. 1946, Compton 18714

(NBG); near Sandberg (-AB), Oct. 1940, Esterhuysen 3460

(BOL); Esterhuysen 3481 (BOL; NBG); Ertjieslandkloof (-AB),

Nov. 1946 Leighton 2277 (K); erroneously as Pillans 2277

(BOL; PRE); Hartebeestkloof, 1 000 m (-AB), Nov. 1974,

Oliver 5169 (K; NBG; PRE; STE). Without locality: Ceres

Wildflower Show Oct. 1936, Compton 6589 (BOL; NBG;

PRE); Oct. 1934, Compton 4933 (BOL.)

E. calycina is probably the showiest of all the

species in the genus having relatively large flowers in

which the corolla, calyx and bracteoles are all bright

pink. They are borne profusely on a compact shrublet.

For this the species is distinct in the genus. Floral

characters which serve to distinguish the species are

the wide-open obconic corolla, aristate anthers and

Fig. 7. — Eremia calycina. Twig, 1,5. I, flower, 18. 2, corolla, > 18. 3, bracteole, X 18. 4, calyx,

ventral view, 18. 5, sepal, inner surface, X 18. 6, stamen, lateral view, X 18. 7, anther, back

view ■< 36. 8, anther, side view 36. 9, gynoecium 18. 10, gynoecium, cut longitudinally, Xl8.

1 1, whorl of leaves, x 18. All drawn from the holotype, Compton 4939 (BOL).

E. G. H. OLIVER

39

erect adpressed leaves. It is probably most closely

related to the previous species, E. peltala.

In his paper in 1935 Compton described four

species. He described E. calycina characterizing it by

the large broad calyx segments, which are indexed

and broadly cucullate at the apex and which exceed

the corolla-tube in length. The latter statement is,

however, not borne out by the type description or

figure (cf. Fig. 7 of the present paper) where the calyx

is just shorter than the corolla-tube. The former

characters could easily apply to one of the other

species he described, E. virgata. This latter species he

claimed, was well distinguished in the genus by its

erect virgate or tufted growth and its very short

adpressed leaves. This, in fact, also applies to E.

calycina. There appeared to be no difference between

the two species.

As both erect, virgate and compact to spreading

shrublets occur with varying degrees of calycine

flowers, the two species were combined under the

more appropriate name, E. calycina. The first name

was also preferred, because the type locality of

E. calycina is known whereas the type of E. virgata

was recorded from the Ceres Wildflower Show.

E. calycina occurs only on the Cold Bokkeveld

Plateau, which has an average altitude of 900 m.

It grows on sandy open flats among short shrubs and

restiad clumps sometimes forming large conspicuous

populations. It flowers from September to November

5. Eremia curvistyla (N.E. Br.) E. G. H. Oliver

comb. nov.

Eremiopsis curvistyla N.E. Br. in FI. Cap. 4,1: 390 (1906).

Syntypes: Worcester Division, Niven s.n. (BOL!; K!); Without

locality: Herb. Salisbury s.n. (BOL!; K !) ; Ward s.n. (K.!).

Lectotype: Niven s.n. (K).

Low, spreading shrublet. Branchlets slender, some-

times long and entwining in surrounding vegetation,

puberulous with simple or glandular hairs, the grey

bark splitting irregularly and flaking off to reveal

red-brown wood, with leaves mostly only on the

young branchlets. Leaves 3-nate, curved spreading to

recurved, crowded, up to 1,7 mm long excluding

the petiole, mostly 1 ,0 mm long, oblong-lanceolate

to ovate, somewhat trigonous, flat above towards the

petiole, convex towards the apex, sparsely pubescent on

upper surface when young becoming roughly scabrid,

otherwise glabrous with an apical gland-tipped hair

and with 3 or 4 on each side which soon become

short setae, sometimes also shortly ciliate; petiole up

to 0,5 mm long, adpressed ciliate. Flowers (2) 3 (4)-

nate on ends of short lateral branchlets, usually

pendulous with 1, 2 or 3 branchlets in a whorl forming

a congested spike-like group; pedicel very short,

about 0, 1-0,2 mm long, glabrous; bracteoles 3,

approximate but spreading, subequal, the median

broadly ovate, acute, 1 x0,6 mm, the laterals broadly

elliptic, obtuse, oblique, all long ciliate with a few

short gland-tipped hairs towards the apex otherwise

glabrous, soft scarious, white becoming reddish,

sometimes with a green leaf-like apex, keel-tipped.

Calyx 4-lobed, divided from two-thirds to three

quarters its length, up to 1,8 mm long, cyathiform

from a narrow base; tube up to 0,5 mm long, quad-

rangular; lobes up to 1, 4x1,3 mm, orbicular to

broadly elliptic, curved and spreading upwards from

the narrow tube, like the bracteoles soft scarious,

white becoming reddish with an acute distinct wide

keel at apex, glabrous or pubescent at apex and

inside the keel, long-ciliate and with many to a few

short glandular hairs towards the apex. Corolla

4-lobed, divided for about a third of its length from

1,7-2, 5 mm long, obconical to campanulate with a

short pouch-like base, sometimes 4-angled with

distinct ridges down the lobes, very sparsely pubescent

on the lower half only, sometimes with a few hairs

up the lobes otherwise glabrous, white; lobes very

broad, obtuse or acute, erect with acute interstices,

smooth or crenulate. Stamens 8 free; filaments

filiform, glabrous; anthers bipartite, 0,4-0, 8 mm long,

included or manifest, haphazardly arranged, narrowly

oblong, dorsally attached near the base, scabrid,

pale brown with darker red-brown backs, aristate;

awns coming off the filament at the point of attach-

ment, about quarter the length of the cell, scabrid;

pore about one-third the length of the cell; pollen

grains single. Ovary 1 (2)-celled, 0,7x0, 6 mm,

pubescent at apex, rarely completely glabrous; when

1-celled with a single, oblique, pendulous ovule,

obliquely ovoid abaxially with the style arising from

the eccentric apex of the ovary and curved upwards;

when 2-celled with a single pendulous ovule in each

cell and apical style; style crooked at the apex

markedly so in the bud, included or exserted and

becoming straighter, glabrous; stigma simple. Fig. 9

A species sprawling in habit, sometimes forming

mats, on dry mountain slopes in sandy rocky places

from Bainskloof northwards to Citrusdal and

Piquetberg.

Cape. — 3218 (Clanwilliam): Piquetberg mountain (-D?),

Oct. 1950, Pillans 10558 (MO). 3219 (Wuppertal): Warmbaths

(-CA), Sept. 1912, Edwards in BOL 14420 (BOL); Turret Peak

near Skoongesig (-CC), Sept. 1962, Esterhuysen 29687 (BOL);

Olifants River Valley, Grootfontein area, 550 m (-CC), Oct.

1972, Oliver 4072 (BM; C; E; G; K; L; LD; MEL; MO; P;

PRE; S; STE; W); Porterville mountains, Kromboskloof,

975 m (-CC), Oct. 1972, Oliver 4091 (E; K; MEL; MO; PRE;

S; STE; W); Graskop above Boboskloof, 970 m (-CC), Oct.

1968, Williams 1330 (NBG; STE). 3319 (Worcester); Twenty-

four Rivers Mountain above Porterville (-AA), Oct. 1949,

Esterhuysen 16082 (BOL; PRE); Twenty-four Rivers valley,

junction of Groot and Kliphuis Rivers, 610 m (-AA), Oct.

1972, Oliver 4100 (E; K; MO; PRE; STE; W); Great Winter-

hoek, 1 524 m (-AA), Jan. 1926, Andreae 1113 (NBG; PRE;

STE); Sneeugat, 1 220 m (-AA), Jan. 1952, Esterhuysen 19817

40

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREMIELLA

Fig. 9. — Eremia curvistyla. 1, flower. 2, corolla. 3, bracteoles: a, median; b. laterals. 4, anther,

front, side, back views. All drawn x20 from the lectotype, Niven s.n. (K). 5, bracteoles:

a, median; b, lateral, both drawn x20 from Hafstrom <6 Lindeberg s.n. (STE). 6, flower.

7, bracteoles: a, median; b, laterals. 8, calyx lobe. 9, gynoecium. 10, ovary, cut longitudinally

to show ovule. All drawn x20 from Oliver 4072 (STE).

E. G. H. OLIVER

41

(BOL; NBG; PRE); Roodesandberg, Tulbagh, 914 m (-AC),

Sept. 1936, Compton 6664 (BOL); Wolwekloof Forest Reserve,

Bainskloof (-CA), Oct. 1946, Barker 4245 (K; NBG; STE);

Bainskloof (-CA), Sept. 1936, Hafstrom & Lindeberg s.n.

(BOL; K; S; STE). Without locality: Niven s.n. (BOL; K);

Herb. Salisb. s.n. (BOL; K); Ward s.n. (K).

On ovary structure E. curvistyla is distinct in the

genus having an oblique 1 -celled ovary and markedly

eccentric curved style. In its sprawling habit it is

similar to E. recurvata, but is easily distinguishable

by its shortly ciliate calyx as opposed to the long

hirsute calyx of the latter.

The species is very similar to Grisebachia parviflora

(see under Excluded Species) which was originally

erroneously described as a species of Eremia. A

comparison with the figure of this species to be

published later in this series will show the remarkable

similarity. The most noticeable difference lies in the

generic character of stamen number — 8 in E. curvistyla

and 4 in G. parviflora. Externally, the large bracteoles

and sepals of E. curvistyla are very different from the

narrower ones of the latter species. The two species

are sympatric with G. parviflora having the wider

distribution range.

This species was first described by N. E. Brown as

a separate monotypic genus, Eremiopsis, characterized

by its unusual ovary, oblique, 1 -celled with an eccentric

markedly curved style. On first examination this

condition suggests an abortive 2-celled ovary. An

investigation of 14 different collections showed this

character to be constant. However, a collection,

Esterhuysen 29687 from the northern Cold Bokke-

veld, when examined was found to have an equal

quantity of 1 -celled oblique ovaries and 2-celled erect

ovaries as in Eremia.

Consideration was given to retaining the species

in a separate monotypic genus. With the above

definite overlap of generic characters it was felt that

separation could not be upheld. The concept of “a

genus of convenience”, which will appear several times

in the treatment of the Ericoideae, could not be

considered in the case of a monotypic genus being

kept separate from an already variable genus.

. 11— Variation in bracteoie size in trenua curvisiyia. i ne seine. u.agia...

ranges in 15 specimens and the map gives the locality of each specimen. The lines A-B, C-L>

and foci X and Y are referred to in the text. 1, Edwards, s.n 2, Herb '-Salisb. s.n 3, Niven s.n.

4, Compton 6664. 5, Esterhuysen 16082. 6, Oliver 4091. 7 Oliver 4100. 8 Esterhuysen 29687.

9, Oliver 4072. 10, Williams 1330. 11, Barker 4245. 12, Andreae 1113. 13, Esterhuysen 19817.

14. Pillans 10558.’ 15, Hafstrom & Lindeberg, s.n.

42

STUDIES IN THE ERICOIDEAE. I. THE GENERA E REM I A AND EREMIELLA

The variation found in E. curvistyla occurs in the

size of the flowers and in the shape and size of the

bracteoles. Fig. 1 1 shows the distribution of bracteole

sizes (length x breadth) of 15 collections. The

median bracteole was included as it is subequal in

nearly all of the collections. The specimens are

numbered in order of increase in size ratios and are

plotted on the map accordingly.

There is a distinct disjunction in bracteole size at

line A-B. The odd ratio is of an unusually developed

median bracteole and can be ignored. Unfortunately

only one of the three specimens is localized, Edwards

s.n. from the Olifants River Mountains above

Warmbaths. Without further exact collections in this

group no taxonomic recognition can as yet be given

to it. In the future, if there are further collections an

assessment of this disjunction can be made.

The remainder of the collections are slightly

separable along line C-D given two foci X & Y of

maximum overlap. This is correlated with a slight

distributional grouping as shown in the map. The

collection Oliver 4072 (No. 9) is anomalous falling

on the wrong side of the line C-D. Pillans 10558

(No. 14) is similarly anomalous.

In his type description N. E. Brown cited three

syntypes, but labelled the sheet of Niven s.n. as the

type. Niven s.n. is now selected as the lectotype.

6. Eremia recurvata Klotzsch in Linnaea 12: 498

(1838); Benth. in D.C., Prodr. 7: 700 (1839); N.E. Br.

in FI. Cap. 4,1: 333 (1905). Type: Cedarbergen,

Drege s.n. (B, holo.|; isos?). Neotype: Cedarberg,

near Ezelbank, Drege 2965 (E, neo.!; BM!; BOL!;

G!; G-DC!; K!; MO!; P!; PRE!; W!).

Shrublets, sprawling or erect up to 30 cm. Branches

rigid, spreading, minutely puberulous with a few

glands or rarely gland-puberulous, with later some

long white hairs admixed which are sometimes hispid,

becoming glabrous and grey, occasionally with

distinct ridges below the leaf-bases when young.

Leaves 3-nate, erect, spreading-recurved to squarrose-

recurved, 2x1 mm, ovate to broadly ovate, fairly

flat at base becoming thick and rounded towards

apex, at first minutely puberulous above becoming

glabrous with some long spreading hairs which may

be gland-tipped, also with subsessile glands, apiculate

with a long gland-tipped hair; petiole adpressed,

shortly ciliate. Flowers 1-10-nate on ends of branches;

pedicels up to 1,5 mm long, pubescent sometimes

with long white hairs as well; bracteoles 3, sub-

approximate to approximate, usually more or less

spreading, usually leaflike, up to 1,7x0, 9 mm, the

median broader than laterals which may be linear-

oblong and acute, all pubescent towards the base,

long-ciliate, often gland-tipped, also with sessile

glands on margins, slightly sulcate. Calyx 4-lobed,

joined for quarter to half its length, up to 2,3 mm

long; lobes up to 0,9 mm broad, narrowly ovate-acute

to ovate-acute, thinly beset with long soft hairs and

with dark sessile to subsessile glands on edge of inner

surface, edged with fine short hairs towards the base

and finely pubescent, sometimes glandular, very

slightly sulcate, forming a cup at the base, often

with 4 main ridges opposite each segment and 4

opposite the interstices. Corolla 4-lobed, to 3,5 mm

long, conical to campanulate, obscurely to markedly

4-angled, glabrous except for thin pubescence near

apex of each lobe and sometimes spread down the

ridge, very noticeable in bud stage, white; lobes very

broadly rounded, erect. Stamens 8, free, included;

filaments up to 1,2 mm long, linear to narrowly

oblong-elliptic, glabrous, transparent but dark at

the apex, tapering into the sigmoid apex below the

anther; anthers up to 0,7 mm long, narrowly ovoid,

narrowed upwards with contiguous separate cells,

bigibbous at the base, minutely scabrid, awned,

golden brown; awns narrowly lanceolate, from

half to equal the length of the cell, ciliate, pale;

pore very small; pollen grains single. Ovary 2-celled,

with a single pendulous ovule in each cell, 0,5 mm

long and broad, broadly ellipsoid to globose-ovoid,

obtuse, very sparsely pubescent to pubescent at the

apex; style filiform, glabrous, included to shortly

exserted; stigma simple. Fig. 12.

Fig. 12. — Eremia recurvata. 1,

flower, X 10. 2, bracteoles:

a, median; b. lateral, both

x20. 3, sepal, inner surface,

x20. 4, stamen, x20. 5,

anther, front, side and rear

views, x20. 6, androecium

and gynoecium, X 10. All

drawn from the neotype,

Drege 2965 (E).

E. G. H. OLIVER

43

A species mostly sprawling in habit occurring in

the drier regions of the south-western Cape between

Ceres and Clanwilliam, found mainly in dry sandy

rocky places.

Cape. — 3219 (Wuppertal); Between Pakhuis and Heuningvlei

(-AA), Dec. 1941, Esterhuysen 7446 (BOL; K); vlakte at

Heuningvlei (-AA), Jan. 1949, Esterhuysen 15012 (BOL);

Northern Cedarberg between Koupoort and Boontjieskloof,

914 m (-AA), Oct. 1945, Esterhuysen 12197 (BOL; NBG; PRE);

Kradadouwsberg, I 500 m (-AA), Dec. 1941, Esterhuysen 7507

(BOL; K); Dec. 1941, Stokoe in SAM 55051 (BM; NBG; PRE;

SAM); Krakadouw Heights, 1 220 m (-AA), Oct. 1923, Pocock

697 (STE); Blaawberg (7-AA/CA), Jan. 1831, Drege s.n. (P);

1 200-1 500 m, Dec., Drege s.n. (SAM); Middelberg Plateau

(-AC), Dec. 1941, Compton 12719 (NBG); Dec. 1939, Ester-

huysen 2464 (BOL; K); Middelberg, 1 500 m (-AC), Dec. 1941,

Esterhuysen 7226 (BOL; K; NBG; NH; PRE); Uitkyk Peak,

1 500 m (-AC), Dec. 1941, Esterhuysen 7367 (BOL; K; PRE);

Cedarberg near Ezelbank, I 220 m (-AC), Dec. 1830, Drege

2965 (BM; BOL; E; G; G-DC; K; MO; P; PRE; W); Sneeu-

berg shale band, 1 676 m (-AC), Dec. 1964, Taylor 6133 (K;

PRE; STE); Matjiesrivier (-AD), Jan. 1944, Wagener 291

(NBG); Gideonskop in S. Cedarberg, 1 220 m (-CB), Nov.

1939, Stokoe 8350 (BOL; K; NBG; NH; PRE); Schurweberg

Peak, 1 370 m (-CD), Jan. 1962, Esterhuysen 29439 (BOL;

STE); Schurweberg above Excelsior, 1 220 m (-CD), Dec. 1961,

Oliver 1606 (PRE; S; STE). 3319 (Worcester): Hartebeestkloof,

S. slopes of Vaalboskloofberg, 1 200 m (-AB), Nov. 1974,

Oliver 5146 (STE). Cold Bokkeveld, mountains near Kleinvlei,

1 670 m (-AB), Jan. 1897, Schlechter 10053 (BM; BOL; E; G;

K; MO; P; PRE; STE; W; Z).

Fig. 13. — Distribution of Eremia recurvata.

This species can be easily recognized by its long-

ciliate calyx, conical often markedly 4-angled corolla,

dilated filaments and anthers with very small pores.

In habit and general appearance it is very similar

to E. curvistyla. It has no closely related species.

E. recurvata does not have much variation. Bentham

described the stamens as mostly 8, sometimes seen

as 6 or 7. This I have not found in any of the material

I examined.

In some specimens from the northern part of the

distribution range e.g. Stokoe in SAM 55051 from

Krakadow and Esterhuysen 7446 from the Pakhuis

Pass area, there is a tendency for the plants to possess

more glands on the branches, leaves, bracteoles and

sepals. On the branches in particular there are short-

stalked glands in addition to the longer gland-tipped

hairs.

The Drege neotype seems to be the only collection

with narrow linear bracteoles; all others have broader

more elliptic leaflike bracteoles, particularly the

median bracteole. Klotzsch in his type description

describes the bracteoles as linear, thus confirming

his examination of the Drege material. However, he

describes the ovary as glabrous. This is somewhat odd

as the Drege material I have dissected is the most

pilose of all the specimens examined.

The holotype in Berlin is no longer extant.

Klotzsch gave only “Cederbergen-Drege” in his type

description. Numerous Drege specimens were dis-

tributed to herbaria, some numbered, some with a

locality in the Cedarberg and some with a locality

which cannot be traced with any certainty (Blaaw-

berg). The material distributed as “Cedarberg prope

Ezelbank” is labelled as Drege 2965 in ten herbaria.

Unfortunately most of the specimens are rather

scanty. The specimen in Edinburgh is better than

most and is chosen as the neotype.

7. Eremia brevifolia Benth. in DC., Prodr. 7: 700

(1839); N.E. Br. in FI. Cap. 4,1: 334 (1905). Syn-

types: Cape, Attaquaskloof, George Division, Masson

57 (BOL!; K!; MEL!); Niven 85 (BM!; G-DC!;

K!). Lectotype: Masson 57 (K).

Erect sparse shrublets to 30 cm. Branches erect,

puberulous with long stiffer often gland-tipped hairs

inbetween. Leaves 4-nate, up to 2,0 mm long, oblong-

ovate to ovate, incurved-spreading with petiole

adpressed, mostly open-backed, subacute, glabrous

but ciliate and edged with sessile glands and one or

two gland-tipped hairs on the lower half; petiole

glabrous, gland-edged. Flowers 3- to 5-nate, terminal

in involucrate heads; pedicel almost none to 0,6 mm

long; bracteoles 3, large, approximate, unequal, the

median larger 1, 7x1,3 mm or 1,3 mm x 1,3 mm,

transversely broadly ovate, the laterals up to 1,3 mm

long, obliquely ovate to elliptic, all acute, pilose in

middle to varying degrees, ciliate and edged with

sessile red glands, keel-tipped, green turning reddish.

Calyx 4-partite, equal to unequal in shape, the

segment opposite bracteoles spathulate, two laterals

obliquely spathulate abaxially, outer sepal narrowly

oblong, otherwise all narrowly oblong, acute, pilose,

gland-edged and slightly ciliate, with or without

keel-tip, green. Corolla 4-lobed, 1,5x1, 7 mm,

ellipsoid to broadly ellipsoid with 8 convolutions in

lower half alternating with stamens, sparsely

puberulous on upper half, white; lobes very broadly

obtuse, erect. Stamens 8, free; filaments 0,7 mm

long, placed in convolutions of corolla, glabrous;

anthers 0,9 mm long, oblong, slightly projecting

below middle with contiguous parallel cells, dorsally

basifixed, manifest to half exserted, bearded on front

edge otherwise glabrous, golden brown, muticous;

pore two-thirds length of the cell; pollen grains

single. Ovary 2-celled, with 1 pendulous ovule per cell,

ovoid, pilose, no nectaries present; style filiform,

glabrous; stigma capitate, from manifest to exserted.

Fig. 14,

44

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREMIELLA

Fig. 14. — Eremia brevifolia. 1,

flower, abaxial view. 2,

flower, adaxial view. 3,

flower with bracteoles and

calyx removed. 4, 3

bracteoles. 5, sepals: a,

abaxial; b and c, laterals; d,

adaxial. 6, anther, front, side

and back views. 7, gynoe-

cium. All drawn from the

lectotype, Masson 57 (K).

8, sepals: a, abaxial; b and c,

laterals; d, adaxial, all drawn

from Oliver 4128 (STE). All

drawn x 10.

Fig. 15— Distribution of Eremia brevifolia.

E. G. H. OLIVER

45

Very localized sparse erect shrublets found only on

the mountains around the old Voortrekker Pass in

Attaquaskloof in the Langeberg Range north-west of

Mossel Bay.

Cape. — 3321 (Ladismith), Attaquaskloof, alpine (-DD), Nov

1773, Masson 57 (BOL; K; MEL) Niven 85 (BM; G-DC; K);

summit of old Voortrekker Pass in Attaquaskloof, 838 m

(-DD), Nov. 1972, Oliver 4128 (BM; BOL; C; E; G; K; MEL;

MO; NBG; P; PRE; S; STE; W).

Eremia brevifolia is easily distinguishable from the

other species in this genus by its capitate involucrate

inflorescences, free narrowly oblong sepals, large

bearded anthers and capitate stigma.

Two distinct forms can be recognized in the

available three collections based on the sepal shape:

Form A: Abaxial sepal spathulate, lateral sepals

obliquely spathulate, adaxial sepal narrowly oblong

( Masson 57).

Form B: Sepals all narrowly oblong ( Niven 85

and Oliver 4128).

All three collections come from the same general

area, Attaquaskloof, with only Oliver 4128 being

an exact locality. Niven 85 matches Oliver 4128 and

most probably came from the same population.

Without any exact information about the spatial

relationship between the two forms A and B no

further comment can be made at present. Only when

a thorough examination of the Attaquaskloof moun-

tains has been made can the correct taxonomic

position of these forms be worked out.

Masson travelled through the Attaquaskloof in

the company of Thunberg in November 1773.

Bentham described Eremia brevifolia in DC. Prodr.

and based his description on two specimens, one in

De Candolle's herbarium, the other belonging to

Lee, both according to Bentham collected by

Masson. The specimen in Geneva is, however, Niven

85, not a Masson collection. Thus Bentham

unknowingly based his species on two syntypes.

It has been stated above that the Masson and

Niven collections can be assigned to two different

forms on the shape of the sepals. Bentham's type

description describes the calyx as “calycis laciniis

oblong-spathulatis”. He could thus have only been

referring to the Masson sheet in Herb. Lee, now at

Kew. This has been selected as the lectotype. The

Niven sheet in Geneva has not been dissected and

examined critically to see whether it matches the

Niven material in the British Museum or Kew.

1 accept that they are duplicates.

There is some confusion about the collectors of

the early specimens as N. E. Brown (1905) queries the

collector of Niven 85. The label on the specimen in

the British Museum was in my opinion written by

Niven as No. 85. The sheets marked No. 57 are in a

different hand and labelled as Fr. Masson, but there

appears to be no guarantee of this as the hand-

writing does not match exactly that on letters in the

Kew Archives. N. E. Brown complicated the issue by

changing an almost identical label on the type of

Grisebachia niveni from Fr. Masson to Niven.

Masson states on the label of the Kew specimen

that the plants are 3 or 4 ft high. Those seen by

myself on the summit of the Voortrekker Pass were

at the most 1 ft (30 cm) high and this was in old

established vegetation that had not been burnt for

some considerable time.

The taxonomic position of Eremia brevifolia is

somewhat obscure. It is placed within the genus

Eremia on a summation of characters, but bears no

relationship to any of the other species included in

that genus. It is the only species in the genus with

4-nate leaves. As has been stated under the phyto-

geographical section, the species is geographically

far removed from the main distribution centre of the

genus in the south-western Cape.

The possibility cannot be ruled out that evolution

of this species has taken place independently of the

rest of the genus Eremia , both having evolved from

ancestral Erica stock by reduction in the cells of the

ovary and number of ovules.

In general appearance, there is a remarkable

similarity between Eremia brevifolia and Eremiella

outeniquae Compton, which occurs on mountain

peaks not far east of Attaquaskloof. Both have 4-nate

similar leaves and flowers in small, terminal, capitate

clusters where the bracteoles are enlarged and larger

than the sepals. Both have a similar habit, although

the latter can form low compact shrublets. Both

grow in similar open dry places at high altitudes in

the Langeberg and Outeniqua Mountains.

However, in the internal floral parts the remarkable

combinations found in Eremiella outeniquae serve

to distinguish them easily. Eremiella having 3 corolla

lobes, 6 stamens and a 3-celled ovary.

EXLUDED SPECIES

Eremia lanata (Klotzsch) Benth. in D.C., Prodr.

6: 700 (1839) based on Hexastemon lanatus Klotzsch

in Linnaea 12: 220 (1838) = Acrostemon xeranthemi-

folius ( Salisb .) E. G. H. Oliver comb, nov., based on

Erica xeranthemi folia Salisb. in Trans. Linn. Soc.

6: 339 (1802).

Eremia parviflora Klotzsch in Linnaea 12: 498

(1838) = Grisebachia parviflora ( Klotzsch ) Druce in

Rep. Bot. Soc. Exch. Club. Br. Isl. 1916: 625 (1917).

Eremia rhodopis H. Bol. in J. Bot. 1894, 239 (1894)

— Erica rhodopis (H. Bol.) //. Bol. in FI. Cap. 4, 1 : 97

(1905).

Eremia tubercularis (Salisb.) Benth. in DC., Prodr.

7: 700 (1839) = Erica tubercularis Salisb. in Trans.

Linn. Soc. 6: 330 (1802): Guth. & Bol. in FI. Cap.

4,1: 97 (1905).

EREMIELLA

Eremiella Compton in J.S. Afr. Bot. 19: 119 (1953).

Type species: E. outeniquae Compton.

The name is the diminutive form of Eremia and

refers to the superficial resemblance to some of the

species in that genus.

Perennial, woody, shrublets erect up to 30 cm or

low and compact about 15 cm high. Leaves 4-nate,

erect or spreading. Flowers 4-nate, in terminal erect or

semipendulous heads. Bracteoles 3, adpressed to the

flower, subequal to unequal, conspicuous and larger

than the sepals, sparsely pubescent and subscarious.

Calyx 4-partite, small compared to the corolla,

sparsely hirsute and subscarious. Corolla 3-lobed,

divided for about quarter of its length, cyathiform,

sparsely hirsute. Stamens 6, free, included. Anthers

bipartite with the cells completely free and shortly

stalked, basifixed, muticous. Pollen grains single.

Ovary 3-celled, with a single, pendulous, apical ovule

46

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREMIELLA

in each cell, broadly ellipsoid and much enlarged in

fruit. Nectariferous disc distinct. Style filiform, far-

exserted, deciduous. Stigma capitellate.

A monotypic genus confined to the Outeniqua and

Tzitzikama Mountains of the Southern Cape.

The genus was described by Compton very recently

in the history of the minor genera of the Ericoideae.

He found when identifying iiis collection from

Ruyterskop near Mossel Bay that he was unable to

place it in any known genus. After summing up the

characters he decided that it was worthy of recognition

as a distinct new monotypic genus.

The possession of 3 bracteoles, a 4-partite calyx,

3-lobed corolla, 6 free stamens with markedly bipartite

anthers and a 3-celled ovary with a single pendulous

ovule in each cell serves to distinguish Eremiella

from all other genera in the Ericoideae.

The most closely related genus is Eremia which

differs in having 4, 2 or 1 -celled ovaries, 8 stamens and

a 4-lobed corolla. To have included this species in

Eremia would have meant a further considerable

emending of the circumscription of that genus.

It was decided to retain Eremiella as a distinct

monotypic genus.

In distribution Eremiella is far removed from the

majority of species of Eremia which occur in the

Western Cape. But the anomalous species, Eremia

brevifolia, grows in an area adjacent to the type

locality on Ruytersberg and is superficially similar to

Eremiella outeniquae.

Eremiella has undoubtedly evolved from some

ancestral Erica stock by way of a reduction in the

number or floral parts, i.e. 4 to 3-lobed corolla,

8 to 6 stamens, 4 to 3-celled ovary and numerous to

one ovule per cell. In this evolutionary pathway Eremia

brevifolia could well have been involved, but resulting

in a combination of characters which classifies the

species as belonging to the genus Eremia.

The single species Eremiella outeniquae forms a

woody shrublet, erect up to 30 cm or low and compact

about 15 cm high growing at high altitudes near the

summits of a few scattered peaks in the Outeniqua

and Tzitzikamma Mountains.

The type locality, Ruytersberg just east of the

Robinson Pass in the Mossel Bay district, was

visited to establish the habitat of the species. Here the

plants were very localized on dry stony slopes facing

south-west and west near the summit. The associated

vegetation was very sparse and consisted almost

entirely of low scattered restiad clumps. The habitat

appeared to be remarkably dry, but no doubt water is

regularly deposited in the area by the south-easterly

and southerly cloudbearing winds in summer.

Eremiella outeniquae Compton in J.S. Afr. Bot

19: 120 (1953). Type: Ruyterskop, Mossel Bay

Compton 21818 (NBG, holo.!; BOL!; K!; PRE!).

Low, compact shrublet up to 15 cm or erect soft

shrublet up to 30 cm. Branches slender, hirsute.

Leaves 4-nate, up to 2 mm long without the petiole,

erect and imbricate to spreading with adpressed

petiole, narrowly ovate-oblong to elliptic-oblong,

acute, semi-openbacked, hirsute with long simple

hairs and sometimes with a few gland-tipped hairs on

the lower edges, ciliate with subsessile glands towards

the apex which ends in a gland-tipped hair; petiole

up to 0,7 mm long ciliate. Flowers terminal, 4-nate,

in closely packed heads which are erect or semi-

pendulous; pedicels from very short up to 0,75 mm

long, glabrous or pubescent; bracteoles 3, subequal

to unequal, approximate and clasping the flower, the

median 1, 5x1,0 mm, elliptic-oblong, the laterals

1 ,5x0,6 mm, narrowly oblong to obovate, all acute,

ciliate and sparsely pubescent on lower half, with a

few dark sessile glands near the apex, keel-tipped,

subscarious, greenish to reddish pink. Calyx 4-partite,

reaching up to the corolla interstices, 1,5x0, 5 mm,

narrowly ovate-oblong to oblong from a broader

base, ciliate and pubescent, with subsessile glands

towards the apex, acute, subscarious, greenish to

reddish pink. Corolla 3-lobed, divided about £ its

length, 2x2 mm, cyathiform, sparsely hirsute to

sometimes almost glabrous with hairs only on the

lobes; lobes broad, obtuse, emarginate, sometimes

ciliate towards the interstices. Stamens 6, free;

filaments filiform, glabrous; anthers included,

bipartite, 0,75 mm long, with the cells obovoid,

completely free and shortly stalked, basifixed, minutely

scabrid, pale brown, muticous; pore about \ the

length of the cell; pollen grains single. Ovary 3-celled,

with a single pendulous apical ovule in each cell,

about 0,5x0, 5 mm, broadly ellipsoid becoming

broadly ovoid in fruit, with a nectariferous disc;

style filiform, far exserted, glabrous, deciduous, up to

2,5 mm long; stigma capitellate. Fig. 16, reproduced

from Compton’s original drawing.

A low compact shrublet confined to the summits of

a few high peaks in the Outeniqua and Tsitsikama

Mountains of the southern Cape.

Cape. — 3322 (Oudtshoorn): Ruyterskop, 1 370 m (-CC),

Nov. 1949, Compton 21818 (BOL; K; NBG; PRE); Nov. 1972,

Oliver 4115 (K; MO; PRE; STE); Jonkersberg, 1 220 m

(-CC), Dec. 1951 Esterhuysen 19387 (BM; BOL; S; STE);

Cradockberg (-CD), Jan. 1940, Zinn in SAM 55896 (SAM;

STE). 3323 (Willowmore): Formosa Peak (-DC), Jan. 1940,

Stokoe 7280 (BOL; STE).

This is the only species recorded in the genus.

In the five collections so far made there is very little

morphological variation. The only noticeable variation

occurs in the leaves, those from the Formosa Peak and

Jonkersberg collections being more erect and imbricate

than in the type which has more widely spaced and

patent leaves.

The four localities in which the species grows occur

in two disjunct areas. The three, Ruytersberg, Jonkers-

berg and Cradockberg, are in the mountain range

north of Mossel Bay and George. Stokoe’s record

from Formosa Peak in the Tsitzikamma Mountains

is 115 km to the east. A search will have to be made

of all the intervening high peaks to establish if this

disjunction is real or due to a lack of records.

The collections all possess mature flowers only.

Those of Zinn and Stokoe collected in January have

considerably enlarged ovaries. A visit to the Ruyters-

berg population in mid-November showed that the

true flowering period must be from about September

to December.

U1TTREKSEL

Hierdie is 'n hersiening van die genus Eremia waarin

7 spesies erkenning geniet en die monotipiese genus

Eremiella bespreek word. Beide genera behoort tot die

Ericaceae- Ericoideae en is endemies in die suidwestelike

deel van die Kaapprovinsie. Die hersiening toon die

noodsaaklikheid om die monotipiese genus Eremiopsis

N.E. Br. onder Eremia te plaas. Hierdie is die eerste

in 'n reeks oor die kleiner genera van die Ericoideae

in Suidelike Afrika.

E. G. H. OLIVER

47

Fig. 16 .—Eremiella outeniquae. 1, twig, x$. 2, flower, abaxial view. 3, flower, side view. 4, flower,

adaxial view. 5, corolla. 6, median bracteole, 7 and 8, lateral bracteoles. 9, 10, 11, 12, sepals.

13, stamen, lateral view. 14, stamen, front view. 15, gynoecium. 16, ovule, 17, fruiting ovary.

18, seed. 19, leaf, abaxial surface. 20, leaf, side view. All x 20, drawn from the holotypc, Compton

21818 (NBG).

\ v

48

STUDIES IN THE ERICOIDEAE. I. THE GENERA EREMIA AND EREMIELLA

REFERENCES

Bentham, G., 1839. Ericaceae. In DC., Prodr. 7: 580-733.

Paris.

Bentham, G., 1876. Ericaceae. In Benth. & Hook, f., Gen. PI.

2: 592. London: Reeve.

Bolus, H., 1905. Erica L. In W. T. Thiselton-Dyer, FI. Cap.

4, I: 96-98. Ashford: Reeve.

Brown, N. E., 1905. Eremia D. Don. In W. T. Thiselton-Dyer,

FI. Cap. 4,1: 332-335. Ashford: Reeve.

Brown, N. E., 1906a. Grisebachia Klotzsch. In W. T. Thiselton-

Dyer, FI. Cap. 4,1: 349. Ashford: Reeve.

Brown, N. E., 1906b. Eremiopsis N.E. Br. In W. T. Thiselton-

Dyer, FI. Cap. 4,1: 390. Ashford: Reeve.

Compton, R. H., 1934. Plantae novitates Africanae. J.S. Afr.

Boi. 1: 144 151.

Compton, R. H., 1953. Plantae novitates Africanae. J.S. Afr.

Bot. 19: 120.

Don, D., 1834. An attempt at a new arrangement of the

Ericaceae. Edin. New Phil. J. 17: 150-160.

Don, G., 1834. Ericaceae. In Gen. Syst. 3: 828. London:

Rivington.

Drude, O., 1897. Ericaceae-Ericoideae. In Engl. & Prantl,

Pflanzenfam. 4,1: 57-65. Leipzig: Engelmann.

Klotzsch, J. F., 1838a. Ericearum, genera et species. Linnaea

12: 211-247.

Klotzsch, J. F„ 1838b. Addenda. I.c. 598-9.

Phillips, E. P., 1926. Ericaceae. In Gen. ed.l, 460. Pretoria:

Division of Botany and Plant Pathology.

Phillips, E. P., 1944. Notes on the minor genera of the

Ericaceae. J.S. Afr. Bot. 10: 69-73.

Rach, L. 1855. Die Ericaceen der Thunberg’schen Sammlung.

Linnaea 26: 789.

Weimarck, H. 1941. Phytogeographical groups, centres and

intervals within the Cape flora. Lunds Univ. Arsskr.

37: 1-143.

Bothalia 12, 1: 49-52 (1976)

Studies in the Ericoideae. II. The new genus Stokoeanthus

E. G. H. OLIVER*

ABSTRACT

The new monotypic genus Stokoeanthus is described, its type and only species being the new species

S. chionophilus E. G. H. Oliver. The position of the genus in the subfamily, its relationships and the reasons

for its establishment are given.

Resume

ETUDES SUR LES ERICOIDEAE. II. LE NOUVEAU GENRE STOKOEANTHUS

Le nouveau genre monotypique Stokoeanthus est decrit , son type et son espece unique etant la nouvelle

espece S. chionophilus E. G. H. Oliver. La position du genre dans la sous-famille, ses relations et les raisons de son

etablissement sont donnees.

Stokoeanthus E. G. H. Oliver, gen. nov. in Erica-

ceae— Ericiodeae ad Blaeria accedens, sed ovario

bicellulari plerumque ovulis tribus pendulis distingui-

tur, ceterum bracteolis tribus calyce 4-partito corolla

4-lobata et staminibus quatuor liberis dignoscenda.

Frutex ad suffrutex. Folia ericoidea. Flores parvi

terminales 2- ad 4-nati in ramulis brevibus lateralibus

aggregati in fasciculos densos versus extrema ramorum ;

bracteolis 3. Calyx 4-partitus ordinatus in seriebus

duabus, sepalis exterioribus interioribus latioribus

sed eidem longitudini. Corolla 4-lobata hypogyna

parva ovoidea calycem superantia. Stamina 4 libera

hypogyna; filamentis gracilibus; antheris bipartitis

minute aristatis manifestis; pollen in tetradis. Ovarium

2-cellulatum plerumque ovulis tribus pendulis in

quoque cellulo interdum 2, 4, 5 vel 6; stylo exserto

filiformi; stigmate infundibuliformi demum peltato

papillis duabus distinctis.

Shrub to shrublet. Leaves ericoid. Flowers small,

terminal, 2- to 4-nate on short lateral branchlets

aggregated into dense clusters at ends of main bran-

ches; bracteoles 3. Calyx 4-partite, arranged in two

series, the outer slightly broader than the inner but

of the same length. Corolla 4-lobed, hypogynous,

small, ovoid, longer than the calyx. Stamens 4, free,

hypogynous; filaments slender; anthers bipartite,

minutely awned, manifest; pollen in tetrads. Ovary

2- celled, with usually 3 pendulous ovules per cell,

sometimes with 2, 4, 5 or 6; style exserted, filiform;

stigma funnel-shaped becoming peltate with two

distinct papillae.

Type species: 5. chionophilus E. G. H. Oliver.

A monotypic genus endemic in the south-western

Cape.

Stokoeanthus chionophilus E. G. H. Oliver sp.

nov.

Frutex erectus virgatus ad 1,2 m altus. Rami

juniores pubescentes demum pilis longis plumosis

admixtis. Folia 3-nata erecta imbricata ad patentia

petiolo adpresso, linearia ad anguste ovata, ad 3,5 mm

longa, glabra interdum ciliato-marginata et caespite

ad apice, acuta sed rotundata. Flores terminales

3- nati interdum 2 vel 4 in ramulis brevibus lateralibus,

aggregati in fasciculos densos versus extrema

ramorum ; pedicello glabro brevissimo, viridi ad rubro ;

bracteolis 3 approximatis adpressis, mediana 1,5 X

0,5 mm anguste ovata, lateralibus 1,25x0,3 mm

anguste oblongis curvatis ad partem exteriorem,

omnibus glabris, ciliato-marginatis, lateralibus plus

magis versus extrema, obtusis cum sepalis albis,

* Botanical Research Unit, P.O. Box 471, Stellenbosch.

carina viridi ad rubicunda. Sepala 4 libera in seriebus

duabus, exteriora 1,75x0,75 mm ovata ad elliptico-

ovata, interiora 1,75x0,6 mm anguste elliptico-

ovata, omnia obtusa glabra ciliato-marginata demum

glandibus sessilibus paucis recava et naviculiformia.

Corolla 4-lobata 2x1,75 mm late elliptico-ovoidea

convoluta distincte 8-plo in parte inferiore sepalis

opposita et inter sepala, glabra alba; lobis latis obtusis

parum patentibus. Stamina 4 libera; filamentis

anguste linearibus glabris sigmoideis infra anthera;

antheris manifestis 1,25x0,75 mm anguste ovatis

scabro-marginatis dorsale basifixus, aureofuscis breve

aristatis; aristis dentatis pallidus laterale patentibus;

poro parvo 0,5 mm longo; pollen in tetradis. Ovarium

2- cellulatum plerumque ovulis tribus pendulis in

quoque cellulo interdum 4 vel 5, rare 2 vel 6, inaequali

ovoideum compressum laterale 0,75 mm longum et

0,5 — 0,75 mm latum glabrum viridi nectariis quatuor

distinctis rubris staminibus alternantibus; stylo exserto

ad 2,5 mm longo filiformi glabro albo; stigmate

infundibuliformi demum peltato, papillis duabus

distinctis viridi ad rubicundo. Fig. 1.

Type. — Cape, 3418 (Simonstown): Somerset Sneeu-

kop, south-facing slopes on shale band south of main

peak, 1480 m, (-BB), 8 Nov. 1973, Oliver 4790

(STE, holotype; BM ; BOL; C; E; G; GRA; K; MEL;

MO; NBG; P; PRE; S; W; Z).

Erect virgate shrub to 1,2 m high. Branches pubes-

cent when young, later with long plumose hairs

admixed. Leaves 3-nate, erect, imbricate to spreading

with the petiole adpressed, linear to narrowly ovate,

up to 3,5 mm long, glabrous, sometimes ciliate-edged

and with an apical tuft, acute but rounded. Flowers

3- nate, occasionally 2 or 4, terminal on short lateral

branchlets, aggregated into dense clusters at the ends

of main branches ; pedicel very short, glabrous, green to

red; bracteoles 3, approximate, adpressed, the median

1,5 x 0,5 mm, narrowly ovate, the laterals 1,25x0,3

mm, narrowly oblong, curved outwards, all glabrous

and ciliate, the laterals more so towards the apex,

obtuse, like the sepals white with greenish to reddish

keeltip. Sepals 4, free, in 2 series, the outer 1,75 X

0,75 mm, ovate to elliptic-ovate, the inner 1,75 X

0,6 mm, narrowly elliptic-ovate, all obtuse, glabrous,

ciliate, sometimes with a few sessile glands, arched

inwards and boatshaped. Corolla 4-lobed, 2 x

1,75 mm, broadly elliptic-ovoid, with 8 distinct

convolutions in the lower half opposite and between

the sepals, glabrous, white; lobes broad, obtuse,

slightly spreading. Stamens 4, free; filaments narrowly

linear, glabrous, sigmoid below the anther; anthers

manifest, 1,25x0,75 mm, narrowly ovate, scabrid-

edged, dorsally ba: fixed, golden brown, shortly

aristate; awns pale toothed, spreading laterally,

50

STUDIES IN THE ERICOIDEAE. II. THE NEW GENUS STOKOEANTHUS

7

Fig.1. — Stokoeanthus chiotio-

philus. 1, flower; 2, corolla;

3, androecium and gynoe-

cium; 4, bracteoles: a,

median; b, laterals; 5, sepals:

a, abaxial; b, lateral, inner

and outer views; 6, anther,

front, side and back views;

7, ovary: a, whole; b, cut

longitudinally; c, cut trans-

versely. All x 10, drawn from

the holotype, Oliver 4790

(STE).

pore small, 0,5 mm long; pollen in tetrads. Ovary 2-

celled, with mostly 3 pendulous ovules per cell,

occasionally 4 or 5 rarely 2 or 6, unequally ovoid,

compressed laterally, 0,75 mm long and 0,5 — 0,75 mm

wide, glabrous, green with 4 distinct red nectaries

alternating with the stamens; style exserted, up to

2,5 mm long, filiform, glabrous, white; stigma funnel-

shaped becoming peltate, with two distinct papillae

on upper surface, green to reddish.

Cape. — 3418 (Simonstown): Somerset Sneeukop, 900 —

1200 m, ( BB), Dec. 1939, Esterhuysen 3542 (BOL); Stokoe

6069 (BOL; PRE); Nov. 1938, Stokoe 7706 (BOL); 1370 m,

Nov. 1973, Oliver 4787 (PRE; STE); Oliver 4787a (PRE,

STE); 1480 m, Nov. 1973, Oliver 4790 (BM; BOL; C; E; GRA;

K; MEL; MO; NBG; P; PRE; S; STE; W; Z); South slopes

above Landdrost Kloof, 1220 m (-BB), Nov. 1971, Esterhuysen

32733 (BOL; PRE; STE); 1370 m, Nov. 1973, Esterhuysen

33311 (BOL; K; NBG; S; STE). Between Somerset Sneeukop

and Landrostkop, (-BB), Nov. 1939, Stokoe in SAM 55446

(NBG; SAM).

This new species is confined to a very small area in

the Hottentots-Holland mountain range above Somer-

set West where it has been collected on only two high

peaks (Fig. 2.)

The first collection was made in November 1938

by that veteran collector of Cape plants, Thomas

Stokoe. The following year it was again collected

by him and also by another well-known collector,

Miss E. Esterhuysen.

Fig. 2. — Distribution of the

genus Stokoeanthus.

E. G. H. OLIVER

51

Stokoe’s material was sent to Prof. R. H. Compton

for naming. He recognized that it constituted an

undescribed species, but was unable to place it in

any genus known to him. He annotated the material

as a possible new genus. The collections then lay in

the ‘incertae’ of the local herbaria.

The unusual feature of the species is found in the

ovary, which is 2-celled with usually 3 pendulous

ovules per cell, rarely 2, 4, 5 or 6. This character is

unique in the subfamily and does not occur to the

author’s present knowledge, in any other genus.

Of the 23 genera at present recognized in the Ericoi-

deae occurring in Southern Africa there are only a

few which have more than 2 ovules per cell i.e.

Erica, Blaeria, Thamnus, Ericinel/a, Philippia, Cocco-

sperma, and Nagelocarpus. The last three genera

possess no bracteoles, unequal calyces and connate

stamens and therefore need not be considered in

any relationship with the species under discussion.

In Ericinel/a the calyx is unequal and there are no

bracteoles, but the stamens are free. This genus is of

the philippioid type and, in addition, it is not part of

the south-western Cape flora. Thamnus which occurs

in the eastern Cape is similarly far removed from the

south-western Cape. It is a unique monotypic genus

having a 1-celled ovary with 4 pendulous ovules

around a free central placenta.

Of the remaining two genera, Erica bears a close

superficial resemblance to the new species. In fact,

the new taxon could easily be mistaken for being a

species of Erica. There are some Erica spp. which

have as few as 1 and 2 ovules per cell in the ovary,

especially those belonging to the section Euryloma,

but none of them has 2 cells. The vast majority of

species in Erica have 8 stamens and only four species

(0,75%) have been recorded with 4 stamens. These

latter, however, have 4-celled ovaries. Thus the combi-

nation of 4 stamens and a 2-celled ovary serves to

distinguish the new taxon from species of Erica.

The possession of 4 stamens is, however, the

only character, although somewhat tenuous, used to

distinguish the genus Blaeria from Erica. This makes

the relationship of the new taxon closer to Blaeria.

But here again the ovary is at variance, as in Blaeria

there are 4 cells in the ovary. The possibility of the

plant being a species of Blaeria with reduction in

ovary parts was examined. This would mean an altera-

tion in the ovary characters for Blaeria , a step which

could only be done on the grounds of a strong relation-

ship between the species concerned. I found no

close resemblance between any of the species of

Blaeria and the new taxon.

The concept of a reduced ovary brought up the

possible relationship with the genera having 2-celled

ovaries with single ovules in each cell. These are

Eremia, Grisebachia, Acrostemon, Simocheilus, Thora-

cosperma, Sympieza, Aniserica, Platycalyx and Coil-

ostigma. After much examination of material from

these genera, I decided that there was no relationship

of any significance between any representatives of

these genera and the new taxon. There is a slight re-

semblance to the genus Eremia. This genus consists

of seven species based mainly in the Ceres district

(see No. 1 of this series). The ovary in Eremia is

either 4-, 2- or 1-celled, but always with a single

ovule per cell and there are 8 stamens.

The relationship of this taxon appears to me to

to be with Erica and Blaeria and to some extent with

Eremia, but it does not fit into any of them as presently

constituted. From Erica it differs in having 4 stamens

and a 2-celled ovary, from Blaeria in having 2 cells

to the ovary and from Eremia in having 4 stamens

and more than 1 ovule per cell. To change the generic

limits of any of these genera to force the inclusion

of the new taxon would, in my opinion, be imprac-

ticable and would cause repercussions in the relation-

ships of and differences between many other genera

of the Ericoideae.

One hesitates to create genera, especially monotypic,

in an already complex and highly variable subfamily,

but after careful consideration I decided to describe

this new taxon as a new monotypic genus and named

it after its discoverer T. P. Stokoe. The combination

of characters serving to characterise the new genus

are the 2-celled ovary with usually 3 pendulous ovules

per cell, rarely 2, 4, 5 or 6, and 3 bracteoles, a 4-

partite calyx, 4-lobed corolla and 4 free stamens.

The position of Stokoeanthus as outlined above

is somewhat dubious. It would appear to belong

to the Erica — Blaeria line of evolution with its reduc-

tion in ovary complement. On the one hand it ap-

proaches closely to Blaeria in the summation of charac-

ters, but not in its outward appearance. The only

species of Blaeria occurring in the same general

area are B. ericoides L., B. dumosa Wendl. and B.

flexuosa Benth. On the other hand, its outward

appearance is very Erica-like.

The variation in ovary arrangement needs further

comment. A large number of flowers was examined

mainly from spirit material of Oliver 4787, 4787a

and 4790 to record ovary variation. The majority of

ovaries contained 3 collateral, subapical, pendulous

ovules in each cell. Occasional cells were found with

4 or 5 ovules and only a few with 2 or 6 ovules. In

the ovaries where there were more than 3 ovules per

cell, the ovules were arranged in two ranks, 3 above

and 2 or 3 below, the lower rank being at the base of

the swollen part of the placenta.

The calyx is described as 2-ranked. This needs

further explanation. The adaxial and abaxial sepals

are slightly broader than the two lateral sepals and

overlap them to a small extent. This condition I

have seen in other genera of the Ericoideae, but

it is not comparable with the philippioid type occur-

ring in Philippia, Salaxis, Coccosperma, Scyphogyne,

Lepterica, Nagelocarpus, Ericinella and Coilostigma.

In these only the abaxial sepal is enlarged laterally

as well as longitudinally, sometimes to a considerable

degree.

Oliver 4787a is worth recording as it possesses

double calyces. Apart from this abnormality the

flowers appear to be normal and functional. This col-

lection was probably a single shrub occurring amongst

normal flowered shrublets, but was not noted at

the time of collecting.

In the area which I visited, the species grew in two

separate populations on the south-facing slopes. The

lower population ( Oliver 4787) consisted ol a small

group of scattered plants growing in a moist-type

fynbos on a boulder-strewn slope and were undoubt-

edly outliers of the much larger population 110 m

higher up the peak.

The large population was confined to the southern

end of the broad shale band of the Table Mountain

Series. This is very prominent at the summit ot the

Hottentots-Holland around Somerset Sneeukop. Here

52

STUDIES IN THE ERICOIDEAE. II. THE NEW GENUS STOKOEANTHUS

the species formed an almost pure community about

1 m high with an understorey of low tussock grass.

Snow renders the shale band particularly con-

spicuous in winter. The large population is then

frequently covered by snow sometimes even as late

as mid-October when the shrubs are in flower. For

this reason the specific epithet, chionophilus, was

chosen.

The dull white to cream colour of the flowers

produces an unattractive shrub even though it flowers

profusely. On being disturbed the shrubs gave off

clouds of pollen. These two factors coupled with the

enlarged stigma gave a strong indication of wind

pollination in the species.

UITTREKSEL

Die nuwe monotiepiese genus Stokoeanthus, sy tipe,

asook die nuwe en enigste spesie, S. chionophilus

E. G. H. Oliver , word beskryf Die posisie van die

genus in die subfamilie, sy verwantskappe en redes vir

die totstandkoming van die genus, word gegee.

Bothalia 12,1: 53-64 (1976)

Notes on African plants

VARIOUS AUTHORS

ASCLEPIADACEAE

NEW SPECIES OF BRACHYSTELMA

The five species described here all appear to have a

restricted distribution and each is based on only one or

two collectings. Efforts have been made over the

years to obtain further material, but without success

and it now seems a forlorn hope for the time being.

There are yet other unnamed specimens in herbaria,

which suggest that the quest for more adequate

material should be intensified.

The lack of a good range of preserved material

applies not only to the five species of Brachystelma

being described here. With few exceptions it applies

throughout the genus. For instance two species are

recorded from the Cape Peninsula, which is accepted

as one of the most thoroughly botanised parts of

southern Africa. B. caudatum (Thunb.) N.E. Br.

( =B . crispum Grah.) was collected at the Cape by

Thunberg soon after 1770, but it was not recorded

positively on the Peninsula until nearly 200 years later,

on a site being cleared for a new housing estate.

B. occidentale Schltr. was collected by R. Schlechter

in 1892 on the slopes above Smitswinkel Bay and it has

not been reported again since.

The rarity of the species of Brachystelma is by no

means their only attraction though. As with the

family Asclepiadaceae as a whole (the orchids of the

Dicotyledons) the coronal structure is of particular

importance in classification. To succeed in giving a

clear picture of the corona in words is a task in itself.

An accurate drawing or photograph is an invaluable

supplement, but even this has its problems, especially

when only dried material is available for dissection.

Considerable skill is required to reveal the three-

dimensional structure of the minute pollinia. The

illustrations accompanying these descriptions will

assist readers to a large degree and I express apprecia-

tion to the artists for them.

Brachystelma delicatum R. A. Dyer sp. nov.,

affine B. nano (Schltr.) N.E. Br., floribus copiosis

fasciculatis, corollae lobis oblongo-ovatis, coronae

lobis exterioribus emarginatis differt.

Herba perennis tuberosa, erecta, sparse ramosa,

5-6 cm alta; tuber subglobosa, 2,5-4 cm diam.;

rami tenues delicati puberuli, pilis decurvis induti:

Folia breviter petiolata, lineari-lanceolata vel-oblonga,

1,5-2, 5 cm longa, 1,5-7 mm lata, subcrassa. Flores

plures fasciculati; pedicelli plusminusve 5 mm longi,

puberuli. Sepala plusminusve lineari-lanceolata, 1,5

mm longa, puberula. Corolla 3,5-4 mm longa,

breviter tubulosa; tubus campanulatus, 1-1,25 mm

longus, 1,25 mm diam.; lobi oblongo-ovati, 2,25-3

mm longi, leviter patentes vel ad apices conniventes.

Corona circiter 1,5 mm alta, basi breviter tubulosa,

biseriata; lobi exteriores suberecti, bilobulati; lobi

interiores circiter 0,5 mm longi, incumbentes.

Pollinia minutissima, circiter 0,25 mm longa, sub-

pyriformia.

Type: Cape, 3326 (Grahamstown), between

Grahamstown and Trappes Valley, 700 m alt. 1965,

D. Hardy 2198 (PRE, holo.).

Perennial dwarf herb with erect stem and globose

tuber 2,5-4 cm diam. Stem erect 5-6 cm tall, slender,

unbranched or sparsely branched from near base,

reddish, minutely puberulous with decurved hairs.

Leaves dark green, somewhat fleshy, linear- to

oblong-lanceolate, keeled, 1 ,5-2,5 cm long, 1 ,5-7 mm

broad, very minutely papillose on margin and keel.

Flowers in fascicles of 4-6 in the axils of the leaves;

pedicels up to about 5 mm long, slender, sparsely and

minutely hairy. Sepals linear-lanceolate or triangular-

lanceolate, about 1,5 mm long, slightly longer than

the corolla-tube, sparsely and minutely pubescent.

Corolla glabrous, 3-4 mm long, with short campanu-

late tube 1 ,25 mm deep and 1 ,25 mm diam. at mouth,

with the sinuses slightly prominent; lobes white with

brown or purple-green tips, oblong-ovate, 2,25-3 mm

long, 1 ,5 mm broad at base, ascending-spreading or

incurved and somewhat thickened at the tips. Corona

lime-green, 1,5 mm high, united at base into a short

campanulate tube 1 mm high, biseriate; outer lobes

divided for about 0,5 mm into 2 obtuse lobules;

inner lobes arising from within the tube, about 0,5 mm

long, incumbent on backs of anthers but not extending

over the staminal column. Pollinia exposed in anther

loculi, more or less pyriform, with translucent upper

1/3 of inner margin, with short delicate caudicles

from near the middle of the inner margin; carrier with

short broad wings on the lower half. Fig. 1.

Fig. 1 . — Brachystelma delicatum , the holotype, life size.

Only two tubers of this species have been recorded.

Mr D. Hardy collected the first on the road from

Grahamstown to Trappes Valley in 1965 and it

54

NOTES ON AFRICAN PLANTS

flowered at the Botanical Research Institute in

December of that year. Col. and Mrs R. D. Bayliss

unearthed a second tuber in the adjacent Coombes

Valley when they were collecting succulents in

August 1968. This came into flower in Grahamstown

on 9th December with the last of the 28 buds opening

partially twelve days later. In this case the corolla-

lobes remained touching at the tips whereas those on

the Hardy specimen opened more or less half way.

The colour of the corolla is white towards the base and

the thickened tips of the lobes are purple-green or

brown. It is a truly dainty species.

Judging from the meagre material available for

comparison and from N. E. Brown’s work on the

genus in FI. Cap. 4,1: 833 (1908), B. delicatum is

nearest allied to B. nanum (Schltr.) N.E. Br. and

B. occidentale Schltr., the former from the O.F.S.

and Transvaal and the latter collected only once on

the Cape Peninsula. The obvious difference is in the

remarkably floriferous habit and there are also

differences in the structure of the coronas.

Cape. — 3326 (Grahamstown), between Grahamstown and

Trappes Valley, Hardy 2198 ; Coombes Valley near Grahams-

town, dry grassland, 700 m, 1968, R.D. & A.W.E. Bayliss 4271.

Brachystelma glenense R. A. Dyer , sp. nov., affine

B. caudato (Thunb.) N.E. Br., corollae tubo extus

glabro, lobis supra breviter pubescentibus, coronae

lobis exterioribus v-formatis distinguitur.

Herba perennis tuberosa e basi ramosa, usque 10 cm

alta; rami recti, glabrescentes vel sparse pubescentes.

Folia breviter petiolata, oblongo-lineata, usque 3 cm

longa, 5 mm lata, supra glabra, infra sparse pilosa.

Flores 2-10, fasciculati; pedicelli 10-15 mm longi,

glabri. Corolla 3,5-4 cm longa, breviter tubulosa,

extus glabra; tubus campanulatus, 4-5 mm longus;

lobi basi ovati, lineari-attenuati, intus infra medium

breviter pilosi. Corona biseriata; lobi exteriores

v-formati, lobulis lateralibus 0,5 mm prolongati;

lobi interiores incumbentes, conniventes.

Type: Orange Free State, 2826 (Brandfort): about

20 km north of Bloemfontein at Glen College of

Agriculture (-CC) on a low ridge, in rooi-grass veld

( Themeda triandra Forsk.) very rare, 3rd March, 1962,

J. W. Mostert 219 (PRE, holo.).

Perennial herb up to about 10 cm tall, branched

from the base, rootstock a tuber, probably up to

about 5 cm in diam.; branches glabrous or thinly and

minutely pubescent. Leaves shortly petiolate, oblong-

linear, up to about 3 cm long and 5 mm broad,

glabrous above, subglabrous below. Flowers 2-10

together from the same flowering eye, fasciculate from

lower nodes, opening more or less together; pedicels

more or less erect, slender, 10-15 mm long, glabrous.

Sepals lanceolate-acuminate, 5 mm long, glabrous.

Corolla 3 , 5-4 cm long, united shortly at base, glabrous

outside; tube campanulate, 4-5 mm long; lobes linear-

attenuate from an ovate base, 3-3 , 5 cm long, shortly

pilose on the inner surface below the middle. Corona

biseriate; outer lobes v-shaped forming pockets

between the inner lobes and with lateral lobules

spreading 0,5 mm from the base of the inner lobes;

inner lobes oblong-obtuse, incumbent and connivent

over the staminal column. Fig. 2.

The known distribution of this species is limited to

its presence in 1952 in an experimental grazing plot on

the Glen College of Agriculture. Mr J. W. Mostert

searched the area for a further specimen both at the

time and in subsequent seasons but with no success.

The rarity of some tuberous-rooted species of Ascle-

piadaceae has been remarked upon before with the

suggestion that it was linked with their edibility. It

must be many years since the tubers were sought

Fig. 2. — Brachystelma glenense, part of the holotype, approxi-

mately life size.

after as a source of food in that area and there has

been ample time for regeneration if it happened

readily. It would be interesting to know what the

limiting factors are.

The name Brachystelma caudatum (Thunb.) N.E.

Br., used for the plant referred to in the comparative

diagnosis above, was placed by N. E. Brown in FI.

Cap. 4,1: 839 (1908) in the synonymy of B. crispum

Grah. A summary of the present priorities is as

follows:

Brachystelma caudatum (Thunb.) N.E. Br. in J.

Linn. Soc. Bot. 17: 169 (1878).

Stapelia caudata Thunb., Prod. 46 (1794).

S. tuberosa Meerburg, PI. Rar. t. 54, Fig. 1 (1789),

not the basionym for, nor equal to Brachystelma

tuberosa R. Br. in Bot. Mag. t. 2343 (1822).

Brachystelma spathulatum Lindl. in Bot. Reg. t.

1113 (1827).

B. crispum Grah. in Ed. Phil. J. 1830, 2: 170 (1830);

N.E. Br. in FI. Cap. 4,1: 839 (1908).

The relationship of B. glenense to B. caudatum must

be close, but here again a detailed assessment is not

possible because of inadequate material. B. glenense

appears to have a more robust inflorescence, the

colour of the flowers differs, the pubescence differs,

even making allowance for considerable variation

in B. caudatum , and apparently there are differences in

the structure of the coronas.

Brachystelma incanum R. A. Dyer, sp. nov.,

herba grosse pubescens, ramis paucis decumbentibus,

foliis obovatis, floribus 1-3, corolla atropurpurea

vel nigrescenti, pedicellis gracilibus, corona late

campanulata, lobis interioribus parvis pulvinatis

distinguitur.

Herba perennis tuberosa, decumbens, grosse pube-

scens, basin versus sparse ramosa, rami usque 10 cm

longi, internodiis 5-10 mm longis. Folia lamina

obovate 1-1,5 cm longa, 1-1,5 lata in petiolum

circiter 1 cm longum sensim contracta. Flores 1-3,

extra axillares producti; pedicelli tenues, 1-1,2 cm

longi. Sepala lineari-lanceolata, 3,5-4 mm longa.

Corolla usque 10 mm diam. 2/3-lobata, extus grosse

VARIOUS AUTHORS

pubescens, intus circum coronam pilis longis albidis

mollibus induta, atropurpurea vel nigrescens; lobi

ovato-lanceolati patentes marginibus recurvatis. Coro-

na late campanulata, 1 , 5 mm diam., lobis exterioribus

saccatis emarginatis, lobis interioribus parvis pulvi-

natis.

Type: Transvaal, 2626 (Klerksdorp), 22 km S.W. of

Lichtenburg, ‘Rooigrasveld’ ( Themeda triandra Forsk.)

about 1 500 m alt. 19/2/1946, J. P. H. Acocks, 12476

(PRE, holo.).

Herb with tuberous root, coarsely hairy. Stem

sparsely branched towards base; branches decumbent,

up to about 10 cm long, shortly pubescent with

curved hairs; internodes 5-10 mm long. Leaves

obovate tapering into a slender petiole about 5 mm

long, 2-2,5 cm long, (including petiole) 1-1,5 cm

broad, pubescent on both surfaces, rarely thinly so on

upper surface, folding upwards. Flowers 1-3, extra-

axillary in succession; pedicels slender 10-12 mm

long pubescent. Sepals linear-lanceolate 3-4 mm

long, somewhat shaggy pubescent. Corolla star-

shaped, up to 1 cm diam., with a very short tube

containing the base of the staminal column, coarsely

pubescent on the outer surface particularly so on the

apical 1/3 of the lobes with recurved margins, dark

maroon to black on upper surface (possibly surface

somewhat uneven) with apical 1/3 of lobes lighter in

colour, with a ring of long (white) hairs around the

corona, otherwise glabrous (the dense hairs on the

under surface may give the impression of marginal

cilia). Corona arising 0,25 m up the staminal column,

broadly campanulate, about 1 ,5 mm diam. and about

equalling the staminal column in height; outer lobes

forming pockets within and emarginate; inner lobes

smaii, cushion-like at the back of the filaments.

(Pollinia absent). Fig 3.

Fig. 3. — Brachystelma incanum, featuring the general hairiness

of the plant, a single flower (1 cm diam.) with long soft

white or whitish hairs surrounding the maroon cam-

panulate corona, in the centre of which is the light coloured

apex of the staminal column, enlarged.

55

It was not my intention to describe this species now

because only one good flower was available for

dissection. But the distinctive appearance of the plant

with hairy, obovate leaves and a ring of long, presum-

ably white, hairs on the corolla surrounding the

corona, giving the whole a rather distinctive hoary

appearance, persuaded me to do so. Mr John Acocks

discovered a single specimen while mapping veld

types in 1946 and it is unlikely that he nor anyone

else will be able to contribute further flowering

material in the near future, a prerequisite for my

part. In addition, with more and more of the ‘Rooi-

grasveld’ habitat coming under the plough, the

survival of the species becomes so much more pre-

carious.

If one must name a species as a close affinity it

would be B. dinteri Schltr., which is similar in its

normally coarsely hairy stems and leaves and flowers.

The species differ in habit, shape of leaves, shape and

pubescence of the corolla-lobes and details of the

corona. B. dinteri was at one time regarded as

restricted to S.W. Africa in its distribution, but in

recent years it has been traced to a few localities in the

northern Transvaal.

Brachystelma remotum R. A. Dyer sp. nov.,

affine B. tuberoso R. Br., caulibus patentibus, floribus

parvis, corollae lobis supra pilis longis indutis, non

ciliatis, corona circiter 2 mm alta, 1 , 5 mm cupulata

distinguitur.

Herba perennis humilis tuberosa, basi ramosa;

rami patentes vel procumbentes, graciles, breviter

pilosi. Folia breviter petiolata, late ovata demum

ovato-lanceolata, usque 1,5 cm longa et lata, supra

dense vel sparse minute pubescentia. Flores 1-2, extra

axillares producti, bracteati; sepala lineari-lanceolata,

usque 3 mm longa, subpilosa. Corolla malvine vel

maculata, 6-8 mm longa, breviter tubulata, extra

glabra; tubus campanulatus, plusminusve 6-7 mm

diam.; lobi triangulati-ovati, 6-7 mm longi, basi 4 mm

lati, marginibus reflexis, supra pilis longis induti.

Corona 2 mm alta, 1,5 mm cupulata, biseriata; lobi

exteriores marsupiiformes, emarginati; lobi interiores

oblongi, incumbentes. Pollinia subglobosa vel pyri-

formia, circiter 0,3 mm longa.

Type: Natal, 2730 (Vryheid): Naauwhoek, near

Utrecht on mountain plateau, shallow soil over rock,

sometimes partially above ground, about 2250 m alt.

Oct. 1962, N. J. Devenish 914 (PRE, holo.).

Perennial herb with tuber and short neck-like stem;

tuber 2,5-4 cm diam., somewhat compressed; stems

branched; branches annual herbaceous, spreading,

3-6 cm long, minutely pilose or subglabrous. Leaves

broadly ovate, sometimes the early ones nearly round,

later ones ovate-lanceolate, obtuse or acute, up to

about 1,5 cm long and broad, minutely pubescent on

upper surface, sometimes thinly so. Flowers 1-2

together, extra axillary, opening in succession.

Sepals linear-lanceolate, up to 3 mm long, with few

long hairs. Corolla mauve or mottled, about 6-8 mm

long, up to 1,8 cm diam. shortly tubular, glabrous

outside; tube broadly campanulate, about 3 mm

long, 6-7 mm wide; lobes triangular-ovate, 6-7 mm

long, 4 mm broad at base, with recurved margins and

conspicuous long white hairs on inner surface.

Corona 2 mm high, united for 1,5 mm in a cup-

shaped tube, biseriate; outer lobes forming emarginate

pockets or pouches; inner lobes arising from inner

margin of tube, incumbent on back of anthers but not

56

NOTES ON AFRICAN PLANTS

meeting over staminal column. Pollinia about 0,3 mm

long, subglobose or subpyriform, attached by short

caudicles to light-coloured carrier. Figs. 4 and 5.

Fig. 4. — Brachystelma r emotum. a, corona: tubular outer

corona, forming deep pockets, with V-shaped margin,

inner corona-lobes arising from within the outer tube,

incumbent on the anthers: b, pollinia and carrier.

Fig. 5. — Brachystelma remotum, plant collected by D. H. Roux,

life size.

Natal. — 2730 (Vryheid): Naauwhoek mountain plateau,

Devenish 914\ between Wakkerstroom and Paulpietersburg on

shale outcrop in grassveld, shallow soil, about 2 000 m,

Oct. 1961, D. H. Roux in PRE, 41785.

The specimens on which this species is based have

been languishing in the National Herbarium for a

number of years. The first tuber was brought to the

National Herbarium by D. H. Roux in 1961 and this

flowered with little delay, wereupon the plant was

photographed and a sketch made of a flower. No notes

were made at the time. A specimen agreeing with the

Roux plant was added to the herbarium collection by

N. J. Devenish in the following year, identified as

B. tuberosum R. Br. It is distinct in several characters,

however, and is possibly more closely allied to

species such as B. pulchellum (Harv.) Schltr. and B.

modestum R. A. Dyer. It is recognized from all these

by the long white hairs on the inner surface of the

corolla-lobes and by the cupular shape of the corona.

Devenish described the flowers as with small mauve

star-shaped corollas and mentioned that the plant

co-exists with Euphorbia clavarioides, Delosperma sp.

and Crassula spp. The corolla-lobes of the Roux

specimen are purple-mottled. The degree of variability

of B. remotum cannot be assessed until more specimens

are studied from its somewhat remote distribution

range on the mountains within the Wakkerstroom,

Utrecht and Paulpietersburg triangle of northern

Natal.

Brachystelma swazicum R. A. Dyer, sp. nov., affine

B. pulchello (Harv.) Schltr., floribus 2-3, breviter

pedunculatis extra axillares productis, coronae lobis

exterioribus bilobulatis, lobulis falcato-curvatis distin-

guitur.

Herba perennis humilis tuberosa, basi sparse

ramosa; rami rubri procumbentes, graciles, pilis

paucis induti. Folia late ovata vel subcordata, usque

2 cm longa, 2 cm lata, petiolata, apiculata, atrovirentia

pilis paucis induta; petiolus 1-1,5 cm longus. Flores

2-3, breviter pedunculati extra axillares producti,

sepala lineari-lanceolata, -±2 mm longa, pilis paucis

induta. Corolla atropurpurea, circiter 4,5 mm longa,

extra glabra, basi breviter tubulosa; tubus late

campanulatus, plusminusve 1,5-2 mm longus, usque

3 mm latus; lobi triangulari-ovati, 3 mm longi, basi

1,5 mm lati. Corona lutea, biseriata; lobi exteriores

1 .5 mm longi, bilobulati, lobulis falcato-curvatis; lobi

interiores oblongi incumbentes. Pollinia subglobosa

vel pyriformia circiter 0,25 mm longa.

Type: Swaziland, 2631 (Mbabane) mountain above

Mbabane, in rock crevices, fully exposed site, 1 400 m

alt., 13 Nov. 1966, R. D. Bayliss 3733 (PRE, holo.).

Perennial herb with rather small dark tuber and 2-3

prostrate branches; branches red, 10 cm or more

long with internodes 1,5-2 cm long. Leaves dark

green, broadly ovate or subcordate, up to 2 cm long

and 2 cm broad, apiculate, petiolate, with a few short

hairs mainly along the margin and on veins; petiole

1-1,5 cm long. Flowers 2-3 from flowering eyes on

rudimentary peduncles outside the leaf axis, develop-

ing successively. Sepals linear-lanceolate, about 2 mm

long, with few long hairs. Corolla dark purple, about

4.5 mm long, shortly tubular, glabrous; tube broadly

campanulate or saucer-shaped, 1 ,5-2 mm long, 3 mm

wide; lobes triangular-ovate, 3 mm long, 1,5 mm

broad at base, glabrous. Corona yellow, biseriate;

outer lobes 1 ,5 mm long, bilobulate with the lobules

falcately curving behind the inner lobes; inner lobes

arising from within the outer lobes, incumbent on the

backs of the anthers, but not meeting over the staminal

column. Pollinia subglobose, 0,25 mm long with

narrow translucent inner margin, sessile or with

very short caudicles, carrier with small narrow oblong

wings. Fig. 6.

Fig. 6. — Brachystelma swazicum. a, corona: tubular base of

outer corona forming deep pockets with 2 falcately

spreading lobules from the margin; inner corona-lobes

arising from within the outer tube, incumbent on the

anthers; b, pollinia with shortly winged carrier.

The distribution of this species is not further

known than the Mountains above Mbabane, Swazi-

land. It was first collected there on 13th Nov. 1966

by Col. Roy Bayliss. Efforts to obtain further material

have so far failed.

l

Since sending the description to press the following

two records have come to light: Transvaal: 2531

VARIOUS AUTHORS

57

(Komatipoort) about 10 km from Baberton on

Havelock Rd., in rock crevices, 22/1/1966, Burtt &

Hilliard 3673 NU; Tienie Louw Nature Reserve,

rocky ridges between boulders, 20/3/1973, Jacobsen

2843 PRE.

At first sight the habit suggests a close relationship

with B. pulchellum (Harv.) Schltr. and its allies, but

the shape of the leaves, the inflorescences and especially

the outer corona-lobes, with falcately curved lobules,

leave no doubt as to its separate specific status.

R. A. Dyer

BLECHNACEAE

FIRST RECORD OF STENOCHLAENA TENUIFOLIA (DESV.) MOORE FOR THE TRANSVAAL

Stenochlaena tenuifolia is a scandent fern, largely o

confined to the east coast of Africa from Kenya in the

north to the eastern Cape Province in the south. All

the South African specimens in PRE were collected

practically within sight of the Indian Ocean. The

species typically occurs in coastal swamp forest,

climbing against rocks and trees to a height of 30 m.

Fig. 7 illustrates the known range of the species in

Southern Africa according to specimens in PRE and

the locality records given by Schelpe in FI. Zamb.

Pteridophyta 242 (1970), which could be traced on

maps.

In 1968 a sterile specimen was collected on the farm

Fourieskraal near Barberton, fully 120 km from the

sea, by Mr M. J. van Biljon, acting on information

supplied by Mr D. A. Jackson of Louws Creek

Timbers. The specimen was taken at the small

Mhlambanyathi Creek near the ruin of the old home-

stead at an altitude of about 550 m. In March 1975

one of us (J.C.S.) visited the site and collected fertile

material. The plants were growing in a small stand

of humid “swamp forest” on leached coarse sandy

alluvium reminiscent of sites on the coastal plain of

Zululand and Mozambique. Important associated

j trees were Syzygium cor datum Hochst., Bridelia

micrantha (Hochst.) Baill., Anthocleista grandiflora

Gilg and Rauvo/Jia caffra Sond. The fern was creeping

over litter on the forest floor and up against tree

trunks.

Because of its close proximity to the ruined home-

stead, it might conceivably have been introduced, but

the plants appear to be growing so naturally in the

forest, that the species is considered to be indigenous

to the area.

Transvaal. — 2531 (Komatipoort): near Barberton (-CB),

Van Biljon sub PRE 38955 ; Scheepers 2401 .

J. C. Scheepers & P. Vorster

ERICACEAE

A NEW SPECIES OF ERICA FROM THE BREDASDORP DISTRICT

Erica occulta E. G. H. Oliver , sp. nov. in genere

singularis propter lobos longos corollae et tubum

brevem, habitum similitudine specierum Aspalathi,

flores occultos et alternationem bractearum folio-

rumque.

Fruticulus dense ramosus, erectus, ramis caespitosis

vel pulvinatis, caulibus principalibus paucis. Rami

tantum vetustiores visibiles, suberosi juniores foliis

omnino tecti, internodis brevissimis, apparenter

glabris. Folia 3(?4)-nata, caespitosa ad extremis

ramulorum, imbricata erecta vel parum incurva ad

9x1 mm cum petiolo, lineares acuta ad obtusa,

complanata versus basim, rotundata versus apicem.

minute sulcata, fere sine distinctione inter laminam

petiolumque, longe ciliata in marginibus et in pagina

abaxiali et versus apice in pagina adaxiali, aliter

glabra; petiolo incolorato piano ciliato; folia aggre-

gata alternantesque bracteis parvis; bractiis breve

triangularibus ad spathulatis ad ovatosubacuminatis

ad 2 mm longis ciliatis longe ciliatis apice aliter

glabris, albis mox fuscescentibus, persistentibus.

Flores singulares terminales in ramulis, subtenti

bracteis; pedicello brevi ad 1 mm longo glabro albo;

bracteolis 3, 1,6-1, 8x1 mm plus minusve oblongis

adpressis ad basim calycis, mediano parum breviore

subacuto, lateralibus obtusis apice obliquo, omnibus

i

58

NOTES ON AFRICAN PLANTS

glabris ciliatisque subviridibus vel albis. Calyx

4-partita; segmentis ad 3,5x2 mm ellipticis ad

obovatis obtusis ad subacutis sulcatis, versus apice

incurvis et carinatus, glabris in dimidio superiore

ciliatis, viridibus. Corolla profunde lobata, 7-8 mm

longa, alba in dimidio inferiore subviridis superiore

mox flavescens, glabra; tubo 1-2 mm longo et ad 2 mm

lato breve cyathiforme; lobis erectis ad 6 mm longis,

oblongis, parum contractis infra medium obtusis.

Stamina 8 libera; filamentis linearibus circa 3,5 mm

longis glabris; antheris inclusis ad 1,9 mm longis

anguste oblongis cum vel sine base gibba in fronte,

terminalibus ad subterminalibus, glabris hinnuleis,

aristis decurrentibus; aristis ad 1 mm longis breve

liberis; poro magno ad 1 ,3 mm longo; granis pollinis

in tetradis formantibus massae pollinis. Ovarium

4-cellulare ovulis numerosis, 1,2x1 mm, late

ovoideum ad ellipsoideum, interdum apice distincto

breve cylindraceo, glabrum, album; stylo 5-6 mm

longo exserto; stigmate initio simp'ice demum minute

peltato processibus quattuor, viridi fuscescenti.

Type: Cape, Bredasdorp District, limestone hills

near Pearly Beach, 7 Sept. 1974, Esterhuysen 33606

(STE, holo, BOL; K; MO; NBG; PRE).'

Shrublet growing from rock crevices on cliffs,

densely branched, erect with main stems adpressed to

the rock faces or caespitose and cushion-like with few

to several main stems. Branches, only the older

visible, gnarled and covered with thick dark brown

irregular cork and the remnants of old leaves, the

younger completely covered with leaves with the

internodes almost non-existant, apparently glabrous.

Leaves 3(?4)-nate crowded together at the ends of

branches with dead leaves persistant on some older

branches, imbricate erect and slightly incurved, up to

9x1 mm including the petiole, linear, acute to

obtuse, flattened towards the base, rounded towards

the apex, inconspicuously sulcate, with almost no

distinction between the lamina and petiole, long-

ciliate on the edges and on the abaxial surface and

towards the apex on the adaxial surface, otherwise

glabrous; petiole colourless, flat, ciliate; leaves

alternating in groups with small bracts which subtend

the flowers; bracts from shortly triangular to spathu-

late to ovate-subacuminate, up to 2 mm long, ciliate,

long ciliate at the apex otherwise glabrous, white soon

turning brown, persistant. Flowers borne singly on the

ends of main branches and subtended by bracts;

pedicel short up to 1 mm long glabrous whitish;

bracteoles 3, 1, 6-1,8 xl mm, more or less oblong,

adpressed to the base of the calyx, the median slightly

shorter subacute, the laterals obtuse with an oblique

apex, all glabrous and ciliate, pale green or whitish.

Calyx 4-partite; segments up to 3,5x2 mm, elliptic

to obovate, obtuse to subacute, sulcate at the apex,

incurved and boat-shaped, glabrous, ciliate in the

upper half, green. Corolla deeply 4-lobed, 7-8 mm

long, white or colourless in the lower half, pale green

in the upper half soon turning yellowish, glabrous;

tube 1-2 mm long and up to 2 mm wide, shortly cup-

shaped; lobes erect, up to 6 mm long, oblong, slightly

contracted } of the way up, obtuse. Stamens 8, free;

filaments elongate linear, about 3 , 5 mm long, more or

t

I

i !'

t

I

I :

It 1

:r

Fig. 8. — Erica occulta. 1, flower,

x 5; 2, corolla, x5; 3, series

of bracts, x 5 ; 4, bracteoles, ; .

x5; 5, sepal, x5; 6, leaf,

abaxial view, x2;7, anther,

side, front and back views,

x 10;9, ovary, x5.Alldrawn i

from Esterhuysen 33606

(STE).

VARIOUS AUTHORS

59

less straight or curved outwards just above the middle,

glabrous, colourless; anthers included, up to 1,9 mm

long, narrowly oblong with or without a gibbous base

in front, terminal to subterminal, glabrous, pale

brown, decurrent-aristate; awns up to 1 mm long,

shortly free with tips slightly spreading; pore large up

to 1 ,3 mm long; pollen grains in tetrads forming pollen

masses. Ovary 4-celled with numerous ovules, l,2x

1 mm, broadly ovoid to ellipsoid, sometimes with a

distinct short cylindric apex up to 0,4x0, 7 mm below

the base of the style, glabrous, white; nectariferous

disc present, colourless; style 5-6 mm long, cylindric,

white turning brown, exserted; stigma at first simple

becoming minutely peltate with four protuberances

arising from its centre, green becoming brown. Fig. 8.

Cape. — 3419 (Caledon): limestone hills near Pearly Beach

I (-DA), 8 Oct. 1972, Esterhuysen 32989 (BOL); 9 Sept. 1974,

Esterhuysen 33606 (BOL; K; MO; NBG; PRE; STE).

This remarkable species recently discovered by Miss

E. Esterhuysen of the Bolus Herbarium is very distinct

in the genus for its tufted cushion-like habit, almost

hidden flowers and much divided corolla.

The latter condition immediately brings to mind a

similar one occuring in Erica lanuginosa Andr., which

is a species of uncertain position in the Ericoideae.

j In E. lanuginosa the corolla lobes are very large, but

! each has a large gibbous base and unique stiff inward

projecting sinus.

Apart from this superficial similarity. Erica occulta

' bears no close relationship to any other species in the

genus and its position in a section of the genus is

uncertain. For convenience, it will probably be best to

place the species in the section Eurystegia with E.

lanuginosa despite the fact that it has such smali

flowers. The general appearance of the flower is also

similar to species such as E. bodkinii and E. grisbrookii,

which occur in the hills of the Caledon and Bredasdorp

districts.

The leaves of E. occulta are extremely crowded

together at the ends of the branches; so much so that

the internodes are almost non-existent. It is thus

difficult to determine the number of leaves per whorl.

It would appear that they are 3-nate. The crowded

tufted leaves and the almost hidden cream flowers give

the species the appearance of some species in the genus

Aspalathus; in fact its true identity is hidden until

one makes a close examination of the material.

The possession of bracts is an unusual feature in the

genus Erica shared only by a few species. Normally

in the genus the flowers alternate with the leaves

and are not subtended by any vegetative organ

and no bracts are present. In E. occulta, however,

there are whorls of bracts, distinct and unlike the

leaves, alternating with the leaves. These bracts occur

at the ends of the branches at the time of flowering

and one of them subtends the single flower. Among

the remaining bracts there are two buds which later

develop into new branches.

In the fresh material examined the pollen remained

aggregated in clumps. This was the case in all of the

flowers examined but may have been an unusual

occurrence. I have not seen this in other species in the

genus.

E. G. H. Oliver

FABACEAE

THE IDENTITY OF CALPURNIA SERICEA HARV.

Harvey, in FI. Cap. 2: 267 (1862), based his descrip-

tion of Calpurnia sericea on a specimen collected by

Von Schlicht in Lesotho. Few, if any, specimens have

been referred to C. sericea subsequently and the

identity of the species has remained in doubt. Phillips,

in Ann. S. Afr. Mus. 9; 479 (1917), cited C. sericea as a

synonym of C. robinioides (DC.) E. Mey. but this was

clearly because he did not have the opportunity of

studying the type material, while Yakovlev in his

treatment of the genus in Nov. Syst. Vyssh. Rast.

8; 181-184 (1971) made no mention of C. sericea at all.

Through the courtesy of Prof. B. Nordenstam,

Curator of the Botany Section of the Museum of

Natural History, Stockholm, the holotype of C.

sericea was received on loan from the Sonder Herba-

rium. The holotype consists of a small flowering twig.

The leaves have 6-8 pairs of leaflets which are densely

appressed-villous on both surfaces but especially

along the midrib on the lower surface. The inflore-

scences are shorter than the leaves and the ovaries are

pubescent on the margins only. Although Harvey

! stated that the holotype was collected by Von Schlicht

in Lesotho (Basutoland), a note on the specimen

written by Sonder says that Harvey was in error in

j assuming that Von Schlicht collected the specimen.

According to Sonder the specimen was collected in

; Lesotho by a missionary whose name was unfortu-

nately unknown to him. As far as is known Von

Schlicht only collected in Namaqualand.

On examining the holotype it was at once apparent

that C. sericea and C. obovata Schinz, in Bull. Herb.

Boiss. 4: 426 (1896), are conspecific and that C.

sericea, being the earlier name, must be adopted for

this species. Yakovlev recognized two varieties within

C. obovata Schinz, namely, var. obovata in which the

ovaries are pubescent on the margins only, and var.

pubescens in which the ovaries are pubescent through-

out. As C. sericea, as it is now to be called, is a fairly

polymorphic species, all of the available material was

examined in an attempt to evaluate the significance of

the degree of pubescence of the ovary in delimiting

varieties within the species.

Examination of the material revealed that the degree

of pubescence of the ovary is not correlated with the

degree of pubescence of the leaflets, with leaflet size,

shape or number, or with geographical distribution.

Codd & Dyer 6263 from the Utrecht district of

northern Natal is a fairly good match of Thode A270

from Kafir Drift-Tweekloof, Utrecht district, northern

Natal, the holotype of var. pubescens, but differs in

having slightly larger leaves. Tinley 628 from Giants

Castle, Natal, which is a good match of Codd & Dyer

6263, differs, however, in that the ovaries are pube-

scent on the margins only. Thus, two specimens with

an almost identical facies key out to two different

varieties. Likewise, specimens with a dissimilar facies

key out to the same variety on the basis of the degree

of pubescence of the ovary. Even among the specimens

which have ovaries pubescent throughout there is

considerable variation in the degree of pubescence

of the leaflet surfaces. In the holotype of var. pube-

scens the upper and lower leaflet surfaces are densely

appressed-villous, in Codd & Dyer 6263 the upper

leaflet surfaces are sparsely appressed-villous and the

lower ±glabrous, while in Marais 1275 from the

eastern Orange Free State the leaflets are ±glabrous

60

NOTES ON AFRICAN PLANTS

throughout. A similar range of variation in the

indumentum of the leaflets is present among specimens

with ovaries pubescent on the margins only.

The degree of pubescence of the ovary has been used

for delimiting subspecies within C. aurea (Ait.) Benth.

but in this species the degree of pubescence of the

ovary is correlated with the degree of pubescence of

the leaflets and with geographical distribution. In

the case of C. sericea, however, the degree of pube-

scence of the ovary is not correlated with any other

morphological character or with geographical distri-

bution. Consequently, it is not intended to uphold the

varieties within C. sericea.

The relevant changes in nomenclature are summa-

rized as follows:

Calpurnia sericea Harv. in FI. Cap. 2: 267 (1862).

Type : Lesotho, collector unknown 82, (S, holo. !).

C. obovata Schinz in Bull. Herb. Boiss. 4: 426 (1896). Type:

Natal, Ingunga, Schlechter 6310 (Z, holo.!). C. obovata var. j |)I

pubescens Yakovlev in Nov. Syst. Vyssh. Rast. 8: 183 (1971). !

Type: Natal, Utrecht district, Kafir Drift-Tweekloof, Thode

A270 (K, holo.!; PRE, iso.!). tffl

C. mucronulata Harms ex Kuntze, Rev. Gen. 3 (2): 54 (1898). i I™

Type: Natal, Klip River district, Van Reenen’s Pass, Kuntze \ |

(K, iso.!).

lie

C. intrusa auct., non (R. Br. ex Ait.f.). E. Mey. sensu stricto. n

J. H. Ross ii

THE TYP1FICATION OF HAEMATOXYLUM D1NTERI

Harms, in Bot. Jahrb. 40: 31 (21 May 1907), based

his description of Caesalpinia dinteri, the basionym of

Haematoxylum dinteri (Harms) Harms in Feddes

Repert. 12: 555 (1913), on Dinter 1169 from Inachab

in South West Africa. Unfortunately the holotype in

the Berlin Herbarium was destroyed during the war,

and efforts to trace an isotype have proved unsuccess-

ful. Although the application of the name H. dinteri

is not in doubt, in the absence of any type material it

seems desirable to select a neotype for this species.

Consequently I now select the specimen Pearson 9717 J

in the National Herbarium, Pretoria, from the j ^

western foothills of the Little Karas Mountains, east of Ik

Holoog Station, Keetmanshoop District, South West

Africa as the neotype of H. dinteri.

J. H. Ross

LILIACEAE

ADDITIONAL NOTES ON THE SOUTH AFRICAN SPECIES OF ASPARAGUS

1. The correct name for A. thunbergianus/. A. Schult.

& J. H. Schult.

In the course of an examination of material in the

Bergius Herbarium in Stockholm, I was shown a

specimen labelled in Bergius’ hand “Asparagus mihi

rubicundus”. This is undoubtedly the type (Fig. 9) of

A. rubicundus Bergius, which was described in his

Descriptiones plantarum ex Capite Bonae Spei (1767).

The specimen was collected at the Cape and communi-

cated to Bergius by Grubb. It cannot be established

whether or not Grubb collected it himself. The

epithet has hitherto not been taken up as the descrip-

tion was considered inadequate and the type had not

been seen.

Baker (Flora Capensis 6: 266; 1896) incorrectly

treated A. rubicundus as a synonym for A. africanus

Lam. var. dependens (Thunb.) Bak. I have retained A.

dependens Thunb. as a synonym for A. africanus, but

do not consider that A. rubicundus should be placed

here. There is no reason to believe that Baker saw

the material of A. rubicundus.

The specimen in Bergius’ Herbarium is a flowering

branch of the species currently known as A. thunber-

gianus J. A. Schult. & J. H. Schult. The correct name

and important synonymy therefore become: —

Asparagus rubicundus Bergius, PI. Cap. Bon. Sp.

88 (1767). Type: Caput Bonae Spei, comm. Grubb

(SBT, holo.!).

A. thunbergianus J. A. Schult. & J. H. Schult., Syst. Veg.

7: 333 (1829); Jessop, in Bothalia 9: 55 (1966).

Asparagopsis dregei Kunth, Enum. PI. 5: 84 (1850). A. thun-

bergii Kunth, Enum. PI. 5: 85 (1850), nom. illegit.

2. Asparagus macowanii Bak.

I formerly recognized two varieties in A. macowanii

Bak. (Bothalia 9: 57; 1966). On the evidence of

herbarium material I believed that the eastern Cape

plants never exceeded 1 m in height, while Natal and

Transvaal material exceeded 1,5 m. An examination

Fig. 9. — Asparagus rubicundus. Holotype in Bergius Herbarium,

Stockholm. ; ^

of many populations in the eastern Cape has shown

that the herbarium record was very incomplete and | !|

that in this area plants may occasionally be up to as k

much as 2 m high. I L

It is, therefore, not possible to recognize A. maco- j

wanii var. zuluensis (N.E. Br.) Jess, as a distinct taxon.

J. P. Jessop

VARIOUS AUTHORS

61

A NEW SPECIES OF ORNITHOGALUM FROM THE NATAL DRAKENSBERG

Ornithogalum aloiforme Oberm. sp. nov. distincta.

Planta minora c. 17 cm alta. Bulbus depresso-

globosus c. 25 mm diam. Folia c. 6 spiram laxam

deposita a basi tubiforma pseudocaulem formant;

lamina linearia acuminata ad 20 cm longa et 3 cm

lata crista excentrica longitudinali glauca mollis

tenuis. Racemus axillaris simplex erectus ad 12 cm

longus floribus compactibis. Perianthii segmenta

albida viridi costata linearia 10 mm longa. Stamina

erecta 8 mm longa; filamenta subulata viridia.

Ovarium obovoideum a basi contractum; stylus

filiformis trisulcatus; stigma trilobatum; ovula multi-

seriata.

Type: Natal, 2929 (Underberg): Giants Castle,

Thabanyama, moist open veld, amongst rocks,

January 1969, c. 3 000 m., Vahrmeijer 1877 (PRE,

holo.). The type flowered at the Botanical Research

Institute, Pretoria during March and April 1975.

Small plant c. 17 cm tall. Bulb depressed-globose, c.

25 mm in diam., fleshy, turning green when exposed

to light. Leaves about 6, persistent, spirally arranged,

close together, the erect basal part tubular, forming

a short pseudo-stem below; lamina spreading,

somewhat laxly recurved, linear-acuminate, up to 20

cm long and 3 cm broad, with a raised longitudinal

ridge to one side, margin smooth, apex acute, dull

glaucous green, soft and thin. Racemes axillary,

simple, c. 12 cm high with the flowers close together

and overlapping; peduncle curved, somewhat flat-

tened; lowest bract largest, subulate. Flower opening

in the morning, closing at night; the perianth-segments

somewhat spreading, linear, about 10 mm long, 4 mm

broad, white with a green dorsal keel. Stamens about

8 mm long, erect; filaments subulate, green; anthers

small, 1 mm, versatile, introrse. Ovary obovoid,

i narrowed towards the base, trisulcate, green; style

white, slender, trisulcate, about as long as ovary;

stigma apical, 3-lobed, minutely papillate; ovules

multiseriate. Figs. 10 and 11.

This species is of great interest because of its habit.

Whereas most bulbous Liliaceae (and all the species

of Ornithogalum that I am aware of) produce annual

leaves and a terminal (or pseudo-terminal) inflor-

escence, this species has the evergreen spirally arranged

leaves forming a pseudo-stem. In their axils appear,

laterally, simple axillary racemes, thus resembling

species of Aloe. In this instance, however, the racemes

are situated in the axils of the leaves and are therefore

lateral, whereas in Aloe they are considered terminal,

being placed on the opposite side of the stem.

In the Liliaceae the ovules are usually biseriately

arranged in the three locules but the above species and

a few others, for instance O. miniatum Jacq., O.

thyrsoides Jacq. etc., have the ovules in a multiseriate

arrangement, probably caused by a doubling of the

placental ridges.

Fig. 10. — Ornithogalum aloiforme. 1, habit, X 1 ; 2, perianth-seg-

ment, x4; 3, stamen, x3; 4, gynoecium, xlO; 5, cross-

section of ovary showing multiseriate ovules, X 6.

Fig. 1 1 . — Raceme of Ornithogalum aloiforme, x i .

A. A. Obermeyer

62

NOTES ON AFRICAN PLANTS

A NEW COMBINATION IN BULBINE

Hermann in his Catalogue of Plants growing in the

Leiden Botanic Garden in 1687, described and

figured on p. 466, t.467, a plant that came from the

Cape of Good Hope belonging to a species that is still

common in the S.W. Cape. He gave it the phrase name

Ornithogalum africanum luteum odoratum foliis cepa-

ceus radice tuberosa. From the good description and

plate it is obvious that it is a Bulbine, for he describes

the barbate filaments and in this genus there is a

section that produces a large rounded tuberous base.

This led Hermann and later Houttuyn and Miller to

retain it in the bulbiferous genus Ornithogalum.

Miller in 1768 bestowed the binomial Ornithogalum

tuberosum on Hermann’s plant in his Gardener’s

Dictionary, ed. 8, No. 10, repeating Hermann’s

description and information that it grows naturally

on the dry rocks at the Cape of Good Hope.

Jacquin in his leones Plantarum Rariorum 2, t .405

(1795) and Coll. 5: 83 (1797) painted and described

this species under the name Anthericum pugioniforme.

Link in his Enumeratio, 1 : 329 (1821) transferred it to

Bulbine, viz. B. pugioniformis (Jacq.) Link. Baker

included it under the latter name in the Flora Capensis

6: 363 (1896), but misapplied the name Ornithogalum

tuberosum Mill., placing O. polyphyllum Jacq. as a

synomym under it on p. 506.

As Ornithogalum tuberosum has not been transferred

to Bulbine the combination is now effected.

Bulbine tuberosa {Mill.) Oberm., comb. nov.

Ornithogalum tuberosum Mill., Gard. Diet. ed. 8, No. 10

(1768). Type: Hermann, Cat. Lugd. t.467.

Fig. 12. — Iconotype of Bulbina tuberosa (Mill.) Oberm., t. 467

in Hermann, Cat. Lugdb. (1768).

A. A. Obermeyer

A NEW SPECIES OF ALOE FROM THE HUMANSDORP DISTRICT

Aloe pictifolia Hardy, sp. nov.

Suffrutex ad 30 cm altus, 18 cm diam. Caules

arcuato-ascendentes, ramosi; rami in rosulam folio-

rum terminantes; foliorum bases persistentes. Folia

conferta, basi amplexicaulia, 12-15 cm longa, 1-2,5

cm lata, sensim attenuata apice acuto vel acuminato,

glauca; supra aliquantum plana, versus apicem

canaliculata, copiose et regulatim albo-maculata;

subtus convex carina prope apicem dentata, albo-

maculata; margo dentibus pungentibus 1 mm longis

4-5 mm remotis armatus. Inflorescentiae 2-4 simul-

taneae, simplices, arcuato-ascendentes; pedunculi 20

cm longi, basi plano-convexi vel complanati, 5-7 mm

lati, 4-5-bracteati. Racemi cylindrato-acuminati,

multo-flori, sublaxi, 14-17 cm longi 3,5-4 cm diam.;

alabastra patentia; fiori nutantes; bracteae late

ovatae, abrupte acutae, 10 mm longae, basi 3-4 mm

latae; pedicelli 11-12 mm longi. Perianthium scarla-

tinum, ad ostium viride cylindrato-trigonum, 15-16

mm longum, prope basin 3-4 mm diam.; basis

rotundata; segrnenta ad basin libera. Antherae

aurantiacae, 1-2 mm exsertae.

Type: Cape, Humansdorp District, Patensie area,

Marais sub PRE 32328 (PRE, holotype).

Plants of shrubby growth up to 30 cm tall. Stems

arcuate-ascending, branched from the base, short or up

to 12 cm long, rosettes dividing and subdividing up to

18 cm across; old leaf bases persistent. Leaves crow-

ded, basally sheathing-amplexicaul, 12-15 cm long,

suberectly spreading, 1-2,5 cm broad at the base.

gradually narrowing to an acuminate apex; upper

surface flat to channelled especially towards the

apex, glaucous, copiously and regularly white-spotted ;

lower surface convex with a toothed keel near the

apex, spotted; margins armed with red-brown, pun-

gent teeth up to 1 mm long, 4—5 mm apart, the inter-

spaces the same colour as the leaf. Inflorescences 2-4

borne simultaneously, simple, arcuate-ascending;

peduncles 20 cm long, base plano-convex to flattened,

5-7 mm broad with 4-5 sterile, many-nerved, ovate

to broadly-ovate bracts, 7-8 mm broad and tapering

to an abrupt, acute point. Racemes cylindric-acumi-

nate, sublaxly many flowered; 14-17 cm long and

3,5-4 cm diam., the buds spreading obliquely to

horizontally, open flowers nutant; floral bracts

broadly ovate, abruptly tapering to an acute point, 10

mm long, 3-4 mm broad at the base; pedicels 11-12

mm long, red-brown. Perianth scarlet red (R.C.S. 1),

greenish at mouth, cylindric-trigonous, 15-16 mm

long, 3-4 mm diam. near the base, base rounded;

outer segments free to the base, 3-nerved, the nerves

obscure for the greater part of their length, turning

greenish at the apex, apices straight, subacute,

scarcely spreading; inner segments free, whitish-

yellow with thin edges, keel colour of the perianth

for four fifths of its length turning green near the

apex, apices more obtuse and slightly greenish on the

inside. Stamens : filaments white, flattened; anthers

orange, exserted by 1-2 mm. Style filiform, stigma

eventually exserted by 1-2 mm long, 1 mm diam., pale

green. Capsule not seen. Figs. 13 and 14.

VARIOUS AUTHORS

63

Fig. 13. — Aloe pictifolia, habit.

Fig. 14. — Aloe pictifolia. One of rosettes with inflorescences.

64

NOTES ON AFRICAN PLANTS

This new species was first collected in 1971 by Mr

G. X. Marais, formerly of Pretoria and now of

Newcastle in Natal, in the Humansdorp District of the

southern Cape. The precise locality has not been

revealed, because it is feared that Aloe enthusiasts

might deplete the natural populations in the veld.

This plant has obviously escaped detection until now,

because of its situation in stony, rugged country.

The species appears to belong to the series Echinatae

(Humiles), the imple inflorescence and cylindric-

trigonous perianth being not unlike those of A.

krapohliana Marl. In general habit and leaf texture it

is, however, very different: the plant is branched

from the base with narrow, copiously and regularly

spotted leaves. The plant has been called A. pictifolia,

because of its “painted leaves”.

D. S. Hardy

MELASTOMATACEAE

A NEW MEMECYLON RECORD FROM TONGALAND

Among the many interesting plants collected by Dr

E. J. Moll in Tongaland in recent years was the

specimen, Moll & Nel 5592, from a small tree up to 4 m

high growing in sand forest 7 km W. of Muzi. The

specimen was clearly referable to the genus Memecylon

but did not match material of either M. natalense

Markgr. or of M. grandiflorum A. & R. Fernandes, the

only two species of the genus previously recorded

from Natal. Comparison of Moll & Nel 5592 with

material in the Kew Herbarium suggested that the

specimen was closest to the recently described M.

sousae A. & R. Fernandes, in Bol. Soc. Brot., Ser.

2,46: 67, t. IV (1972), from Mozambique. The speci-

men differed from Gomes e Sousa 4380, the isotype

of M. sousae in the Kew Herbarium, however, in

having consistently smaller leaves. The largest leaves

on Moll & Nel 5592 were 3, 5x1,7 cm and these

were significantly smaller than those in typical M.

sousae. A small flowering twig detached from Moll &

Nel 5592 was sent to Prof. A. Fernandes for comment.

He kindly confirmed that the specimen was close to

M. sousae but added that if the leaves were consis-

tently small there was a possibility that the specimen

may represent an undescribed species.

In an attempt to clarify the situation Dr Moll was

requested to collect more material and, in particular,

to try and establish the range of variation in leaf

size within a population and even on a single plant.

Unfortunately Dr Moll was unable to locate the

plant on his next visit to Tongaland but the following

year (1972) another population was found 6 km W.

of Muzi and the collections Moll & Muller 5690a,

5690b were made. The leaves on Moll & Muller

5690a (NH) are mostly the same size as those on

Moll & Nel 5592. However, on Moll & Muller 5690b

(NH), which was collected from the same plant as

5690a, the leaves are significantly larger and range in

size from 3, 5-5, 6x2, 1-3, 8 cm. These larger leaves

on Moll & Muller 5690b match those of typical M.

sousae and this specimen was therefore referred to M.

sousae. As Moll & Muller 5690a was collected off the

same plant, it too, together with Moll & Nel 5592, were

also referred to M. sousae. The range of variation in

leaf size within M. sousae is therefore greater than

initially realized and indicated by material from

Mozambique.

J. H. Ross

MESEMBRYANTHEMACEAE

NOMENCLATURE OF THE

While checking some literature it was noticed that

the genus Amphibolia was not validly published as

Bolus (1965) did not cite the type species (Article 37),

but Herre’s account on ‘The Genera of the Mesem-

bryanthemaceae’ in 1972 can be construed as a

validation of the generic name. However, all the

species are invalid as they were described, or their

combinations made, before the genus was validly

published (Article 43). Also, in the case of Amphibolia

hallii and A. littlewoodii an incomplete reference to the

basionym was given. The nomenclature of Amphibolia

is briefly reviewed.

Amphibolia L. Bol. ex Herre, Gen. Mesembryanth.

70 (1971); L. Bol. in J. S. Afr. Bot. 31: 169 (1965);

Jacobsen, Sukk. Lex. 358 (1970). Type species: A.

hallii (L. Bol.) L. Bol. ex Toelken & Jessop.

A. hallii (L. Bol.) L. Bol. ex Toelken & Jessop, comb,

nov. L. Bol. in J. S. Afr. Bot. 31 : 169 (1965); Jacobsen,

Sukk. Lex. 358 (1970), nom. inval.

Stoeberia hallii L. Bol. in J. S. Afr. Bot. 26: 161

(1960). Type: Cape, Karoo Poort, H. Hall 1741

(BOL, holo.).

A. gydouwensis ( L . Bol.) L. Bol. ex Toelken &

Jessop, comb. nov. L. Bol. in J. S. Afr. Bot. 33: 306

(1967); Jacobsen, Sukk. Lex. 358 (1970), nom. inval.

GENUS AMPHIBOLIA

Lampranthus gydouwensis L. Bol. in J. S. Afr. Bot.

29: 13 (1963). Type: Cape, Gydouw, Leipoldt 4801

(BOL, holo.).

A. littlewoodii ( L . Bol.) L. Bol. ex Toelken & Jessop,

comb. nov. L. Bol. in J. S. Afr. Bot. 31: 170 (1965);

Jacobsen, Sukk. Lex. 358 (1970), nom. inval.

Stoeberia littlewoodii L. Bol. in J. S. Afr. Bot. 26:

162 (1960). Type: Cape, 24 km east of Karoo Poort

on the road to Touws River, Littlewood in Karoo

Gardens 522/59 (BOL, holo.).

A. maritima L. Bol. ex Toelken & Jessop L. Bol.

in J. S. Afr. Bot. 31: 169 (1965); Jacobsen, Sukk.

Lex. 359 (1970), nom. inval. Type: Cape, near

Vredendal, H. Hall 2885 (BOL, holo.).

A. stayneri L. Bol. ex Toelken & Jessop, L. Bol. in

J. S. Afr. Bot. 32: 126(1966): Jacobsen, Sukk. Lex. 359

(1970), nom. inval. Type: Cape, between Matroosberg

and Theronsberg Pass, Stayner in Karoo Gardens

258/65 (BOL, holo.).

The authors wish to acknowledge the assistance of

Dr R. K. Brummitt, Royal Botanic Gardens, Kew,

with whom the problem was extensively discussed.

H. R. Tolken & J. P. Jessop

Bolhalia 12,1: 65-109 (1976)

A procedure for standardizing comparative leaf anatomy in the

Poaceae. I. The leaf-blade as viewed in transverse section

R.P. ELLIS*

ABSTRACT

Descriptive “keys”, including definitions and diagrams, for standardizing and simplifying the description of

grass leaf structure as seen in transverse section are given. Over 500 characters are included with the possibility

for expansion to 999. Notes on variation and taxonomic importance of the characters are also included.

Resume

UN PROCEDE POUR STANDARDISER L'ANATOMIE COMPAREE DE LA FEUILLE DE POACEAE. I.

LE LIMBE VU EN SECTION TRAN SVERS ALE

Des “ clefs' ’ descriptives incluant des definitions et des diagrammes pour standardiser et simplifier la descrip-

tion de la structure des feuilles de graminees vues en section transversale sont donnees. Plus de 500 caracteres sont

inclus avec la possibility d' expansion jusqu a 999. Des notes sur la variation et T importance taxonomique des

caracteres sont egalement fournies.

INTRODUCTION

Anatomical investigations of the grass leaf-blade

have long provided valuable taxonomic information.

In fact, nowadays, it is generally accepted that

anatomical details, especially of the leaf-blade and

embryo, when used in conjunction with a wide

spectrum of other diagnostic characters, are an

essential ingredient of any satisfactory treatment of

grass taxonomy. Furthermore, in the Poaceae

(=Gramineae), with their highly specialized and

reduced flowers, very fine morphological distinctions

are often necessary to define differences between

taxa. Anatomical data is, therefore, regarded as

being of undoubted importance in the jigsaw of

complete systematic evidence in this numerically large

and important family.

The importance of anatomy in agrostological

studies has resulted in the rapid accumulation of an

extensive body of literature with attendant problems

of lack of uniformity with definitions and descriptions.

Valuable data is, therefore, commonly not applicable

to the family as a whole and comparisons cannot be

drawn with any degree of assurance. This problem

was greatly ameliorated by the publication in 1960

of Anatomy of the Monocotyledons. 1. Gramineae

by C. R. Metcalfe and the present paper is a further

attempt to stabilize this terminology, and, at the

same time to present a system whereby description

and comparison of grass leaf anatomy will be

simplified and standardized.

DESCRIPTIVE KEYS

In an attempt to achieve the necessary uniformity,

descriptive “keys” have been compiled for use as a

framework for anatomical descriptions of the grass

leaf-blade as viewed in transverse section. The “keys”,

which incorporate both definitions and diagrams,

have been designed to enable the user to standardize

descriptions and the hierarchical layout has been

chosen to facilitate the speed and ease at which

^Botanical Research Institute, Department of Agricultural

Technical Services, Private Bag XI 01, Pretoria.

complete, comparative descriptions can be compiled

Anatomical characters, and all other information

considered to be of diagnostic or taxonomic impor-

tance, and gathered from an extensive survey of

the relevant literature, have been included. The keys

should, therefore, prove adequate for all tribes of the

Poaceae except for the Bambuseae where additional

data on the fusoid cells must be included to satis-

factorily distinguish between certain genera.

The hierarchical tabulation of the characters has

been used to expedite their use but they do not in any

other way conform to any acknowledged key format

or design. However, if a statement, at any level of the

hierarchy, does not apply to the specimen being

examined, no subsequent statements or characters of

a lower rank are relevant. The user, therefore, merely

proceeds to the next character of equal rank or

indentation, and, if applicable, works inwards noting

all the relevant numbered end points before working

back outwards until the same level or rank as that of

the originally chosen statement is reached. Thus, the

“keys” are not dichotomous or true indented keys,

but by using this type of format all the possible

characters are recorded in a constant descriptive

sequence. Each character is assigned a constant

number and the recording of these numbers effects a

saving in time and space. Furthermore, this system

enables easy conversion to edge punched and feature

cards as well as for electronic data processing by

computer. In addition, it is ensured that all possible

structures are rapidly and routinely noted in a

rational sequence which simplifies compilation of

descriptions of the various taxa. By employing a

standard sequence significant differences become more

readily evident.

In order for the standardization of terminology and

descriptions to be effective it is essential that com-

parative material be examined. Therefore, in this

study, all descriptions refer to transverse sections

taken at a point about halfway between the blade

apex and the ligule of mature basal leaves. Flag

leaves on flowering culms were avoided where possible.

By standardizing on the material studied in this

way intraspecific differences may be assessed.

66 A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

I. OUTLINE OF THE LAMINA IN TRANSVERSE SECTION

Outline usually determined by the shape of the abaxial surface. Includes the entire transverse section of the leaf blade at a

point about halfway between the leaf apex and the base of the lamina. Subdivided into leaves that can be or are relatively

flattened and permanently infolded, acicular leaves.

OUTLINE OF THE LAMINA OF OPEN LEAVES:

Includes all leaves which are not permanently infolded to such a degree that the internal structure is altered. These leaves

are normally open or expanded under favourable environmental conditions but exhibit various degrees of infolding or

inrolling under conditions of water stress. Projecting and grooved midribs and keels must be considered in relation to the

overall shape.

DESCRIPTION:

Expanded; commonly flattened; a line connecting both margins and the median bundle straight:

Nature of blade; ribs and furrows not considered:

Flat, even or straight 101*

Undulating gently 102*

Distinctly wavy or undulating:

Corrugated leaf i.e. waves rounded 103*

Pleated leaf i.e. waves pointed or angled 104*

V-shaped; two halves of the lamina folded toward each other on either side of the midrib i.e. a line connecting both

margins with the median bundle angled at the median bundle; carinate:

Angle formed by the two arms of the lamina at the midrib:

Narrow; closed V i.e. less than 45° to each other 105*

Standard V i.e. between 45° and 90° to each other 106*

Broad, open V i.e. more than 90° to each other 107*

Wide, very open V i.e. almost 180°; presence of projecting keel gives appearance of V-shape 108*

Shape of arms of the lamina:

Straight 109*

Outwardly bowed; concave 110*

Outwardly curving; convex Ill*

Outline heart-shaped 112*

Distinctly wavy or undulating:

Corrugated leaf i.e. waves rounded 113*

Pleated leaf i.e. waves pointed or angled; plicate 114*

Irregularly wavy or bent 115*

Symmetry of two halves of the lamina:

Symmetrical on either side of the median vascular bundle 116

Two halves assymmetrical on either side of the median vascular bundle 117*

U-shaped; two halves of the lamina curved upwards on either side of the midrib or with a prominent rounded keel i.e.

no definite angle formed with the midrib; canaliculate:

Shape of the U formed:

Broad, wide i.e. horizontally elongated 118*

Tall, narrow i.e. vertically elongated 119*

Base and sides same length i.e. equidimensional 120*

Inrolled; lamina rolled inwards towards the adaxial surface:

Nature of inrolling:

Convolute; inrolled from ONE margin only; margins wrapped around each other 121*

Involute; inrolled from BOTH margins 122*

Degree of inrolling:

Tightly inrolled 123

Loosely inrolled 124

DESCRIPTION CONTINUED. Measurements taken by estimation utilizing the known diameter of the field of view of the

objective where the thickness or width of the leaf almost fill this field of view. This standardisation is for the Zeiss

Standard RA microscope fitted with X 10 Kpl W eyepieces.

Width of one half of the lamina i.e. from blade margin to midrib or median vascular bundle if midrib indistinguishable:

XI 6 Objective:

Less than half the field of view at x 160 i.e. less than 0,56 mm wide 125

Between half and the full field of view at xl60 i.e. from 0,56 mm - 1 ,125 mm wide 126

X2,5 Objective:

Less than half the field of view at x25 i.e. 1,125 mm - 3,6 mm wide 127

Between half and the full field of view at x25 i.e. from 3,6 mm - 7,2 mm wide 128

Up to twice the field of view at x25 i.e. from 7,2 mm - 14,4 mm wide 129

More than twice the field of view at x25 i.e. more than 14,4 mm wide 130

Thickness of lamina taken at a point about halfway between the median bundle and the margin. Depth of largest rib in

central area of lamina measured:

X40 Objextive:

Less than one quarter of the field of view at x400 i.e. less than 0,1 12 mm thick 131

Between one quarter and half the field of view at x400 i.e. from 0,1 12 mm - 0,225 mm thick 132

Between half and three quarters of the field of view at x400 i.e. from 0,225 mm - 0,336 mm thick 133

Between three quarters and the full field of view at x400 i.e. from 0,366 mm - 0,450 mm thick 134

More than the full field of view at x400 i.e. more than 0,450 mm thick 135

R. P. ELLIS

67

OUTLINE OF OPEN LEAVES

120

121

122

68

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

1. OUTLINE OF THE LAMINA IN TRANSVERSE SECTION CONTINUED

OUTLINE OF THE LAMINA OF PERMANENTLY INFOLDED LEAVES:

Includes only leaves that are permanently infolded to the extent that the internal structure is altered. Often includes

acicular, setaceous, filiform and junciform leaves. The entire section is considered under low magnification with the abaxial

surface determining the outline.

DESCRIPTION:

Shape, outline; as formed by the abaxial surface:

Reduced V-shaped; two halves of the lamina folded toward each other on either side of the median bundle i.e. a

definite angle formed:

Arms straight 150*

Arms angled; often hexagonal 151*

Reduced U-shaped; two halves of the lamina curved upwards on either side of the median bundle i.e. no definite

angle formed:

Arms vertical or parallel to one another 152*

Arms rounded or outwardly bowed:

Elliptical; forming an incomplete oval or ellipse 153*

Rounded; forming an incomplete circle 154*

Triangular; base broad on either side of the median bundle 155*

Circular ox Round in transverse section:

Hollow cylinder; margins not fused; adaxial surface not reduced; horseshoe-shaped 156*

Solid cylinder; margins fused or nearly so:

Adaxial surface reduced to a minute groove 157*

Adaxial surface not distinguishable; radially symmetrical; terete 158*

Number of Vascular Bundles present in section; includes median bundle or midrib if present:

3 vascular bundles in T/S 159

5 vascular bundles in T/S 160

7 vascular bundles in T/S 161

9 vascular bundles in T/S 162

1 1 vascular bundles in T/S 163

13 vascular bundles in T/S 164

15 vascular bundles in T/S 165

17 or more vascular bundles in T/S 166

Adaxial channel bounded by the arms of the lamina; may be open or closed at the leaf margin:

Vertical channel sides; parallel to each other:

Width of the channel; compared with the blade thickness at a point midway between median bundle and margin:

Wider than the leaf thickness 167*

Nearly the same as the leaf thickness 168*

Narrower than the leaf thickness 169*

Narrow cleft; less than half the leaf thickness 170*

Depth of the channel; compared with the blade thickness at the median vascular bundle:

Shallower than the leaf thickness 171*

Nearly the same as the leaf thickness 172*

Deeper than the leaf thickness 173*

Deep; more than twice the leaf thickness 174*

Non-vertical channel sides:

Rounded channel 175*

Triangular channel 176*

Channel reduced to small groove 177*

No channel present 178*

DESCRIPTION CONTINUED. Measurements taken by estimation utilizing the known diameter of the field of view of the

objective where the thickness or width of the leaf almost fill this field of view. This standardisation is for the Zeiss

Standard RA microscope fitted with X10 Kpl W eyepieces.

Width of one half of the lamina i.e. from blade margin to midrib or median vascular bundle if midrib indistinguishable:

XI 6 Objective

Less than half the field of view at xl60 i.e. less than 0,56 mm wide 179

Between half and the full field of view at xl60 i.e. from 0,56 mm - 1,125 mm wide 180

X2,5 Objective:

Less than half the field of view at x25 i.e. 1 ,125 mm - 3,6 mm wide 181

Between half and the full field of view at x25 i.e. from 3,6 mm - 7,2 mm wide 182

More than the full field of view at x25 i.e. more than 7,2 mm wide 183

Thickness of lamina taken at a point about halfway between the median bundle and the margin. Depth of largest rib in

central area of lamina measured:

X40 Objective:

Less than one quarter of the field of view at x400 i.e. less than 0,1 12 mm thick 184

Between one quarter and half the field of view at x400 i.e. from 0,1 12 mm - 0,225 mm thick 185

Between half and three quarters of the field of view at x400 i.e. from 0,225 mm - 0,336 mm thick 186

Between three quarters and the full field of view at x400 i.e. from 0,336 mm - 0,450 mm thick 187

More than the full field of view at x400 i.e. more than 0,450 mm thick 188

R. P. ELLIS

69

OUTLINE OF PERMANENTLY INFOLDED LEAVES

176

177

178

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

2. AD AXIAL AND ABAXIAL LONGITUDINAL RIBS AND FURROWS

Includes all longitudinal ribs or ridges as well as the associated furrows or grooves commonly found on the adaxial leaf

surface but sometimes on the abaxial surface as well. If ribs are present there must be corresponding furrows and vice versa.

If furrows are not found between the ribs but in any other situation they are termed grooves, e.g. the grooves present on

some ribs. Ribs and the associated vascular bundles form regular associations and patterns. The furrows and the adaxial

channel of acicular leaves can be distinguished by the fact that the channel is the area bounded by the permanently

infolded lamina and the furrows lie between the ribs which may also be in the channel.

DESCRIPTION:

No ribs or furrows present on either surface i.e. lamina surface straight or only slightly undulating with no regular

pattern associated with the vascular bundles 201*

Adaxial ribs and furrows present:

Depth of adaxial furrows in comparison to the leaf thickness i.e. the depth of the larger ribs in the central region of

the lamina between the margin and the median vascular bundle regarded as leaf thickness:

Slight, shallow furrows i.e. less than a quarter of the leaf thickness 202*

Medium furrows i.e. a quarter to one half the leaf thickness 203*

Deep funows i.e. more than one half the leaf thickness 204*

Shape of adaxial furrows:

Wide, open furrow i.e. obtuse angle formed by furrow sides at base 205*

Narrow furrow i.e. sides of furrow almost vertical:

In form of cleft 206*

Base fairly broad but sides steep 207*

Distribution of adaxial furrows; important for acicular leaves:

Furrows between all vascular bundles 208*

Furrows between 1st and 2nd order vascular bundles i.e. present over 3rd order vascular bundles 209*

Furrows between 1st order vascular bundles i.e. present over 3rd and 2nd order vascular bundles 210*

Furrow on either side of the median vascular bundle only 211*

Two funows on either side of median vascular bundle only 212*

Deepest furrows on either side of the median vascular bundle 213*

Shape of adaxial ribs as seen in T/S; all types present must be included:

Rounded, obtuse ribs i.e. apex rounded:

Situated over the vascular bundles 214*

Situated between the vascular bundles (usually occupied by air spaces) 215*

Flat-topped, square ribs i.e. apex flattened:

Sides rounded with flat top 216*

Sides angled with flat top 217*

Trangular ribs i.e. apex pointed:

One vascular bundle in each rib 218*

Three vascular bundles in each rib 219*

Grooved at apex 220*

Massive ribs i.e. very large usually with bases narrow in relation to upper parts of rib:

Flattened apices 221*

Rounded apices 222*

Distribution of different ribs in association with the vascular bundles:

Ribs present over all vascular bundles:

Similar ribs over all vascular bundles 223*

Ribs over 1st and 2nd order vascular bundles larger than those over the 3rd order vascular bundles 224*

Ribs over 1st order vascular bundles with flattened tops and those over the 3rd order bundles triangular . . . 225*

Ribs not associated with all vascular bundles:

Ribs present only over 1st and 2nd order vascular bundles 226*

Ribs present only over 1st order vascular bundles 227*

Ribs present over midrib only 228*

Abaxial ribs and furrows present:

No abaxial rib development:

Surface smooth or with few undulations not regularly associated with the vascular bundles 229*

Grooves present on abaxial surface:

Opposite larger vascular bundles 230*

Between vascular bundles; often shallow 231*

Groove on either side of midrib 232*

Ribs present on abaxial surface:

Size of abaxial ribs:

Taller than the adaxial ribs 233*

Same size as adaxial ribs i.e. section resembles a string of beads; moniliform 234*

Smaller than adaxial ribs 235*

Slight undulation associated with the vascular bundles 236*

Distribution of abaxial ribs:

Present opposite 1st order vascular bundles only 237*

Present opposite 1st and 2nd order vascular bundles only 238*

Present opposite all vascular bundles 239*

Composition of abaxial ribs:

Composed of sclerenchyma:

In form of rounded caps 240*

In form of flat-topped caps 241*

Composed of girder or strand of sclerenchyma in contact with the epidermis 242*

RIBS AND FURROWS

R. P. ELLIS

71

241

242

72 A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE PnArp*c

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION ACEAE.

3. MEDIAN VASCULAR BUNDLE, MIDRIB AND KEEL STRUCTURE

ttSed"! mkMh If b5L the.,presen“ 01 absence »f Parenchyma associated with the median vascular bundle It is

oTnchLT^ n, ,rr I!3" bundl,! 1S soUtaIy' alfootarally distinct and without associated parenchyma and a he f

pMenchyma or bullrform cells are associated with the median bundle or bundles. Thus one or many vascular bundled f

to “eea, «f oa^ch™ d°eS "0t 7?*^ ,n **•»*!» “1- «*■*> « is“nsS

Thp u 1 ^ nchyma is present) unless there is a marked thickening in relation to the rest of thp iomi

DESCRE^ION^aSCU 31 bUndl6S ^ 3 ked iS’ ” 3 gCneral W3y’ r°Ughly Pr°P°rtionaI t0 size of the keel.

Not p°Ssible to determine from preparation i.e. entire width of blade not sectioned

“stiTuSlc y Plesent i'e-entile °' m0re **“" half ** blade motioned and no siructuraliy disiinct midrib fa 3°'

Ire^inTOn^icinius:*' tbs,’"8uishable from* other first order vascular bundles; no associated parenchyma developed* be! 3°2*

Projection of midrib:

Projects abaxially:

Sclerenchyma causes projection

Projection due to position or size of bundle 303*

Projection inconspicuous or marked; not projecting abaxially only: 304*

Blade expanded or slightly inrolled

Lamina V-shaped but no parenchyma at angle 305 *

Keel developed!31 colouiless ‘ ZS^nXT be devWopfd bu> "° in association with ih'e midrib' i i

development ^ydToccur: ' bUlhf°rm ““S aSS0Ciated with ,he media" ba"<Ue; sclerenchyma

Number of vascular bundles comprising keel; in V-shaned leaves if th^ ;<■ ... ,

median bundle is considered to constitute the keel:' ** ’ 6 1 n° thlckening in the keel area the

One vascular bundle comprising the keel:

Runif^Im!nf r0,Unde,d ceUs surrounding or immediately adaxial to the median bundle ,AO*

Bulhform cells in adaxial epidermis above median bundle- 308

Leaf expanded, flat or slightly inrolled . . .

Leaf V-shaped : 309*

-v ^

Bullrform cells and round^paren^hymVceUs^esent^3'^ parencbyma 312*

bei"8 3 SU“ adaxia' 3rd order bundle ' i i i i i i $1

Adaxial groove present

No adaxial groove present 315*

abaxial 311(1 oneadaxial amphi^ bundie ' • • • • : : : : : : : : : : 33l76*

All vascular bundles abaxially arranged:

Median bundle indistinguishable from other 1st order bundles-

One 1st order bundle and smaller bundles comprise keel

Three 1st order bundles and other smaller bundles comprise keel 318

Media^^ keel. . : . : .' : .* : : : : : : : : : : : III

v

^TJbS:^ « “d <■ <ba onhe keel; usually htdudK .purposed

Shape of keel in transverse section: 323*

Not really distinct from leaf outline

V-shaped, pointed or inverted triangular keel: 324*

Leaf V-shaped with V-shaped keel

Leaf V-shaped with inverted triangle-shaped keel 325 *

R n,^LeXPanded Wth blverted triangular or pointed keel 326*

R°unded or semicircular keel; leaf not necessarily U-shaped: 327*

U-shaped kee ; shghtly thicker than rest of lamina

U-shaped keel; much thicker than rest of lamina . 328*

RounHpH F TmiC1!rCUlar keel Le- adaxkd ^de of keel fiai * 329*

Rounded keel with raised, flattened adaxial side 330*

Rounded keel with single, central adaxial groove 331*

Rounded kee with distinct adaxial rib developed 332*

Massive j IT? ™merous adax*l ribs and furrows' 333*

« JJ? T U'shaped keel; often with air-spaces 334*

Sclerenchyma associated with the keel: 335*

Adaxial sclerenchyma:

Strands in subepidermal layers

Strands fused forming a hypodermal band 336*

Abaxial sclerenchyma: 337*

Most bundles with abaxial strands

Most bundles with abaxial girders .... 338*

A it °r SOlitary bundle with anchor-shaped girder ’ 339*

breaking do^: mCOrporated in the keel; definite lacunae present' or' enlarged parenchyma ^ of ke'ei ^

Une air-space in keel

Two air-spaces in keel 341*

Four or more air-spaces in keel 342*

. . . . 343*

o

midribs and keels

321

322

323

R P. ELLIS

73

74 A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

4. VASCULAR BUNDLE ARRANGEMENT

Usually studied under low magnification. Only half the width of the lamina, from the margin to and including the median

vascular bundle considered. When the section is incomplete and the median bundle is indistinguishable only those

characters that are possible to determine are considered. In certain cases where the 2nd order vascular bundles are not

clearly distinguishable from the 1st order bundles they are (Considered as 1st order bundles for the purposes of

arrangement.

ARRANGEMENT:

Total number of 1st ORDER bundles in half the width of the lamina; includes the median vascular bundle:

Not possible to determine; entire blade not sectioned 401

1 1st order vascular bundle in blade section i.e. median bundle 402

2 1st order vascular bundles in half lamina i.e. 3 in entire blade 403

3 1st order vascular bundles in half lamina i.e. 5 in entire blade 404

4 1st order vascular bundles in half lamina i.e. 7 in entire blade 405

5 1st order vascular bundles in half lamina i.e. 9 in entire blade 406

6 1st order vascular bundles in half lamina i.e. 1 1 in entire blade 407

7 1st order vascular bundles in half lamina i.e. 13 in entire blade 408

8 1st order vascular bundles in half lamina i.e. 15 in entire blade 409

9 1st order vascular bundles in half lamina i.e. 17 in entire blade 410

1 0 or more 1 st order vascular bundles in half lamina 411

Arrangement and Alternation of different orders of vascular bundle along width of blade; central part of lamina

between margin and midrib studied:

Variable from median bundle to margin; no regular pattern discernible:

Progressively fewer 1st and more 3rd order bundles towards the margin 412

Progressively more 1st and fewer 2nd and 3rd order bundles toward the margin 413

Arrangement near margin differs from remainder 414

Regular arrangement from median bundle to margin:

Number of -3rd order bundles between consecutive 2nd or 1st order bundles at a position halfway to margin:

No 3rd order bundles between consecutive larger bundles 415

1 3rd order bundle between consecutive larger bundles 416

2 3rd order bundles between consecutive larger bundles 417

3 3rd order bundles between consecutive larger bundles 418

4 3rd order bundles between consecutive larger bundles 419

5 3rd order bundles between consecutive larger bundles 420

6 3rd order bundles between consecutive larger bundles 421

7 3rd order bundles between consecutive larger bundles 422

8 3rd order bundles between consecutive larger bundles 423

9 3rd order bundles between consecutive larger bundles 424

10 or more 3rd order bundles between consecutive larger bundles 425

Number of 2nd order bundles between consecutive 1st order bundles:

No 2nd order bundles between consecutive 1st order bundles 426

1 2nd order bundle between consecutive 1st order bundles 427

2 2nd order bundles between consecutive 1st order bundles 428

3 2nd order bundles between consecutive 1st order bundles 429

4 2nd order bundles between consecutive 1st order bundles 430

5 or more 2nd order bundles between consecutive 1st order bundles 431

Position of vascular bundles in blade; vertical situation in the section about halfway between median bundle and

margin:

Same level of positioning for all bundles of different orders:

All bundles situated in the centre of the blade 432*

All bundles situated closer to the abaxial surface 433*

All bundles situated closer to the adaxial surface 434*

Different levels of positioning for bundles of different orders:

1st order bundles central and 3rd order bundles abaxial 435*

3rd order bundles central and 1st order bundles displaced adaxially in ribs 436*

All bundles arranged irregularly and inconsistently 437*

Bundles of all orders abaxial and an adaxial superposed, often amphivasal bundle in each rib 438*

R. P. ELLIS

75

VASCULAR BUNDLE ARRANGEMENT

436

437

431

76

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

5. VASCULAR BUNDLE DESCRIPTION

Vascular bundles are considered to consist solely of the xylem and phloem tissue. For the purposes of describing the shape

of the vascular bundles the bundle sheaths are not included as being part of the bundles. Thus, when a bundle is described

as being circular or angular in outline, these terms refer only to the vascular tissue and exclude the inner or single bundle

sheath.

DESCRIPTION:

Third order vascular bundles; usually very small bundles often with xylem and phloem indistinguishable and consisting

of only a few lignified cells and a few phloem elements; when not obviously smaller than basic type bundles often

distinguishable by the absence of sclerenchyma strands and/or the presence of bulhform cell groups adaxially:

No third order bundles present in section 501

Shape of third order bundles in section:

Rounded in outline; usually with many small parenchyma sheath cells:

Circular in outline 502*

Elliptical; vertically elongated 503*

Angular in outline:

Square-shape or pentagonal i.e. surrounded by a sheath of 4 or 5 large parenchyma cells 504*

Hexagonal or octagonal i.e. surrounded by 6 or more relatively small parenchyma cells 505*

Triangular in outline 506*

Vertically elongated; tall and narrow 507*

Nature of vascular tissue of third order bundles:

Xylem and phloem groups distinguishable 508*

Vascular tissue consists of only a few vascular strands 509*

Second order vascular bundles; xylem and phloem easily distinguishable; bundles usually fairly large, often of similar

size to the first order bundles; no conspicuously large metaxylem vessels or lysigenous cavities present; sclerenchyma

arrangement usually the same as for first order bundles:

No second order bundles present in section 510

Shape of second order bundles in section:

Rounded in outline; usually with many small parenchyma sheath cells:

Circular in outline 511*

Elliptical; vertically elongated 512*

Angular in outline; parenchyma sheath of relatively few large cells; inner mestome sheath may also be present:

Square-shaped or pentagonal 513*

Hexagonal or octagonal 5 14*

Triangular in shape; often rather large 5 15*

Tall and narrow in outline; vertically elongated; bundles with straight vertical sides 5 16*

First order or Basic type vascular bundles; large metaxylem vessel present on either side of protoxylem elements;

lysigenous cavity commonly present; associated with sclerenchyma girders or strands:

Shape of first order bundles in section:

Circular or round in outline 517*

Elliptical; vertically elongated 518*

Egg-shaped; broadest side adaxial 519*

Relationship of phloem to vascular fibres:

Phloem adjoins the inner or parenchyma sheath 520*

Phloem completely surrounded by thick-walled fibres 521*

Phloem divided by intrusion of small fibres resulting in sclerosed phloem 522*

Nature of lysigenous cavity and protoxylem:

Lysigenous cavity and enlarged protoxylem vessel present 523*

Lysigenous cavity but no protoxylem vessel present 524*

Enlarged protoxylem vessel present but no lysigenous cavity 525*

No lysigenous cavity or protoxylem vessel present 526*

Size of metaxylem vessels in relation to parenchyma sheath cells in T/S:

Narrow vessels i.e. parenchyma sheath cells wider than vessels 527

Wide vessels i.e. vessels with width equal to or slightly greater than parenchyma sheath cells 528

Very wide vessels i.e. width of vessels very much more than that of parenchyma sheath cells 529

Shape of metaxylem vessels as seen in T/S:

Angular in T/S 530

Circular in T/S 531

Thickening of walls of metaxylem vessels as seen in T/S:

Unthickened; secondary wall barely visible 532

Slightly thickened; intermediate between unthickened and distinctly thickened state 533

Distinctly thickened; inner secondary wall distinct 534

R. P. ELLIS

77

78 A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

6. VASCULAR BUNDLE SHEATHS

Includes the single or double sheath completely or partially surrounding each vascular bundle and adjoining the mesophyll.

May also include adaxial or abaxial extensions of parenchyma cells which are considered to be part of the bundle sheath

and not part of the system of colourless parenchyma. The different orders of vascular bundles may be surrounded by

sheaths of different structure and thus the sheaths associated with the different orders of vascular bundles are considered

individually.

ORDERS OF VASCULAR BUNDLE present in section; bundle sheaths of each order of vascular bundle present

completely described before continuing with the descriptions of the sheaths of the other orders of vascular bundle present:

Third order vascular bundle sheaths; describe fully: 601

Second order vascular bundle sheaths; describe fully: 602

First order vascular bundle sheaths; describe fully: 603

DESCRIPTION. A complete description is given separately for each order of vascular bundle present:

No bundle sheaths surrounding the vascular bundles 604*

Outer or Single bundle sheath; parenchyma sheath; includes vascular bundles with single and those with double sheaths:

Shape of bundle sheath:

Sheath round, circular 605*

Sheath elliptical, vertically elongated 606*

Sheath triangular with adaxial apex 607*

Sheath tall and narrow with vertical sides 608*

Sheath horse-shoe shaped 609*

Extent of bundle sheath:

Sheath complete; completely surrounding the bundle 610*

Sheath incomplete due to interruption of sclerenchyma girders of various sizes:

Adaxial interruption:

Slight interruption caused by a narrow girder of 1 - 3 fibres wide 611*

Wide interruption caused by broad girder of more than 3 fibres wide or by colourless parenchyma .... 612*

Abaxial interruption:

Slight interruption caused by a narrow girder of 1 — 3 fibres wide 613*

Wide interruption caused by broad girder of more than 3 fibres wide or by colourless parenchyma .... 614*

Sheath reduced to two lateral strips at the sides of the phloem; adaxial interruption not due to sclerenchyma

girders 615*

Extensions of the bundle sheath; comprised of parenchyma cells associated with the sheaths and not part of the

bulliform groups; extend to adjacent sclerenchyma girders or strands or to the epidermis:

No extensions of the bundle sheath present 616*

Nature of the extensions:

Adaxial extension of the bundle sheath:

Narrow extension :

Uni-seriate; consisting of one column of cells 617*

Bi-seriate; consisting of two columns of cells 618*

Broad extension; tri- or multi-seriate 619*

Abaxial extension of the bundle sheath:

Narrow extension ; uni- or bi-seriate 620*

Broad extension; tri- or multi-seriate 621*

Distinctive cells similar to those of the parenchyma sheath found scattered in the mesophyll:

In groups of 2 - 6 with vascular tissue lacking 622*

Solitary 623*

Composition of the extensions:

Consist of large, thin-walled colourless cells; the same size or bigger than sheath cells 624*

Consist of large, thin-walled cells with chloroplasts;the same size or bigger than sheath cells 625*

Consist of relatively small, thin-walled colourless cells; smaller than the sheath cells 626*

Consist of thickened parenchyma cells 627*

Consist of colourless cells gradually decreasing in size as walls increase in thickness and eventually merge into

the sclerenchyma strand 628*

Consist of parenchyma cells on either side of sclerenchyma strands or girders 629*

Extent of the extensions of the bundle sheath:

Extend to and in contact with the epidermis or an inconspicuous sclerenchyma strand (1-4 fibres) 630*

Extend to conspicuous sclerenchyma strand 631*

Extend to sclerenchyma strand almost extending to the bundle sheath 632*

Extend to colourless subepidermal parenchyma 633*

Extend into mesophyll tissue 634*

Length of extension:

1- 2 cells long or deep 635

2- 5 cells long or deep 636

More than 5 cells long or deep 637

Number of cells comprising the bundle sheath:

4 parenchyma cells comprise the sheath 638

5 parenchyma cells comprise the sheath 639

6 parenchyma cells comprise the sheath 640

7 parenchyma cells comprise the sheath 641

8 parenchyma cells comprise the sheath 642

9 parenchyma cells comprise the sheath 643

1 0 parenchyma cells comprise the sheath 644

1 1 parenchyma cells comprise the sheath 645

12 parenchyma cells comprise the sheath 646

1 3 parenchyma cells comprise the sheath 647

14 parenchyma cells comprise the sheath 648

15 or more parenchyma cells comprise the sheath 649

CONTINUE ON FOLLOWING PAGE AND THEN RETURN AND DESCRIBE SHEATHS OF ANY OTHER ORDERS OF

VASCULAR BUNDLES THAT ARE PRESENT.

R. P. ELLIS

VASCULAR BUNDLE SHEATHS

80

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

6. VASCULAR BUNDLE SHEATHS CONTINUED

DESCRIPTION CONTINUED. A complete description is given separately for each order of vascular bundle present:

Structure of the parenchyma sheath cells; : cells of each order of vascular bundle considered separately and all

cell types included:

Not well differentiated from the chlorenchyma cells 650

Distinct from chlorenchyma cells:

Shape of cells comprising sheath; include all shapes relevant for each order of vascular bundle:

Inflated, round, circular 651*

Radial walls straight; tangential walls inflated 652*

Radial and outer tangential walls straight; inner tangential wall inflated 653*

Radial and inner tangential walls straight; outer tangential wall inflated; fan-shaped 654*

Cells elliptical; elongated; often rather irregular 655*

Arrangement of cells of different shapes comprising the bundle sheath:

All cells similar in shape 656

Irregular; cells of different shapes comprise sheath 657

Two enlarged elongated cells abaxiaUy situated:

Adjoining girder of sclerenchyma 658*

Normal parenchyma sheath cells between the elongated cells 659*

Cells at adaxial side of sheath elongated vertically; elliptical 660*

Sheaths of adjacent bundles continous with each other; Triodia type 661*

Size of cells comprising sheath:

Cells of sheath similar in size:

Large and inflated; generally larger than mesophyll cells; conspicuous 662*

Not markedly larger than the mesophyll cells; conspicuous 663*

Cells smaller than the mesophyll cells; inconspicuous 664*

Cells smaller than the inner bundle sheath cells 665 *

Cells small; resemble cells of a mestome sheath i.e. thickened walls often especially the inner tangential

walls; parenchyma sheath apparently absent 666*

Cells of sheath of various sizes:

Cells of different sizes comprise sheath; irregular 667*

Gradation in size with largest cells adaxially situated 668*

Gradation in size with largest cells abaxially situated 669*

Gradation with smallest cells in the centre; abaxial and adaxial cells larger 670*

Gradation with largest cells in the centre on each side 671*

Wall structure of bundle sheath cells:

Walls thin; no secondary thickening 672*

Walls slightly thickened; secondary cell wall not distinct 673*

Walls distinctly thickened; secondary wall distinct 674*

Inner tangential and radial walls thickened 675*

Chloroplast structure of the bundle sheath cells:

Translucent; sheath cells without chloroplasts 676

With small chloroplasts; not distinct from chloroplasts of the chlorenchyma:

Numbers of chloroplasts comparable to the mesophyll cells 677*

Numbers of chloroplasts fewer than in mesophyll cells 678*

With large, specialised chloroplasts:

Chloroplasts fill entire cell lumen 679*

Chloroplasts concentrated near the outer tangential wall 680*

Chloroplasts concentrated near the inner tangential wall 681*

Chloroplasts oentrally situated 682*

Inner or Mestome bundle sheath; endodermis; only applicable when two sheaths are present; often difficult to

distinguish from fibrous ground tissue of the vascular bundle; in contact with the metaxylem i.e. if metaxylem vessels in

contact with parenchyma sheath then there is no inner bundle sheath:

Extent of sheath;

No inner sheath present 683*

Sheath complete; completely surrounding the xylem and phloem 684*

Sheath incomplete due to interruptions of sclerenchyma girders:

Adaxial interruption 685*

Abaxial interruption 686*

Sheath reduced; intermediate type; interruptions not due to sclerenchyma girders:

Abaxial; opposite phloem only 687*

Adaxial; opposite xylem only 688*

Structure of cells of inner bundle sheath in region adjacent to phloem; cells of each order of vascular bundle

considered separately:

Relatively large with inner tangential and radial walls thickened 689*

Small, with inner tangential and radial walls thickened; U-shaped thickenings 690

Small, with uniformly thickened walls:

Walls heavily thickened; lumen small '691

Walls not conspicuously thickened 692

Larger than outer bundle sheath; parenchymatous; with chloroplasts 693

Adaxially situated cells larger than lateral cells of the sheath 694*

N.B. SHEATH OF EACH ORDER OF VASCULAR BUNDLE PRESENT IN SECTION DESCRIBED COMPLETELY

BEFORE CONTINUING WITH SCLERENCHYMA OF THE LEAF.

R. P. ELLIS

81

VASCULAR BUNDLE SHEATHS

82

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

1. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

7. SCLERENCHYMA OF LEAF

Includes all thickened fibrous tissue in the leaf, except that constituting part of the vascular tissue i.e. xylem and phloem

fibres and thickened parenchyma. The sclerenchyma can be sub-divided into the circumvascular sclerenchyma surrounding,

in contact with, or opposite the vascular bundles in the form of girders or strands, sclerenchyma found in the intercostal

areas between successive bundles, sclerenchyma of the leaf margin and sclerenchyma of acicular leaves. Sclerenchyma

associated with the midribs or keels included under midribs and keels.

SCLERENCHYMA STRANDS AND GIRDERS

Circumvascular sclerenchyma surrounding, in contact with, or opposite the bundles; refers only to .longitudinal

subepidermal sclerenchyma following the course above and below the vascular bundles. Complete description is given of

the circumvascular sclerenchyma associated with each of the different orders of vascular bundle present in the leaf.

ORDERS OF VASCULAR BUNDLES present in leaf section; sclerenchyma associated with each order of vascular bundle

present in section considered separately and each is completely described before commencing with other orders that might

be present:

Sclerenchyma associated with third order vascular bundles 701

Sclerenchyma associated with second order vascular bundles 702

Sclerenchyma associated with first order vascular bundles 703

DESCRIPTION

AD AXIAL sclerenchyma; sclerenchyma found above the vascular bundles; associated with the bundles:

No adaxial sclerenchyma present 704*

Strand present; sclerenchymatous; not in contact with the bundle sheath; separated by parenchyma or mesophyll

from sheath:

Minute strand consisting of only a few subepidermal fibres 705*

Small strand; inconspicuous; epidermal cells unaltered over the strand:

Shallow, forming a strip; only 2 - 4 fibres deep; subepidermal 706*

Narrow; only 2-4 fibres wide 707*

As deep as wide as seen in T/S 708*

Well-developed strand; conspicuous; epidermal cells over strand usually small and thick -walled:

As deep as wide as seen in T/S 709*

Wider than deep as seen in T/S; in form of band:

Arched; follows the shape of the adaxial rib 710*

Straight, horizontal band 711*

Girder present; sclerenchymatous; in contact with, or interrupts the bundle sheath:

Very small girder; inconspicuous; epidermal cells usually unaltered over girder 712*

Small girder; epidermal cells over the girder usually small and thick -walled:

Small thin subepidermal strip 713*

Narrow; deeper than wide 714*

As deep as wide as seen in T/S; equidimensional 715*

Well-developed girder; conspicuous; epidermal cells over girder small and thickened:

Relatively wide and deep band; as wide or wider than the vascular bundle:

Arched; follows the shape of the adaxial rib 716*

Straight; horizontal band 717*

Narrowing towards bundle 718*

Relatively narrow and deep girder; narrower than the vascular bundle 719*

T-shaped; horizontal band connected to bundle or bundle sheath by a stem:

Stem short; shorter than horizontal cross-piece 720*

Stem relatively long; as long or longer than horizontal cross-piece 721*

Inversely anchor-shaped; arched band follows shape of adaxial rib:

Stem short and sturdy; more than tri-seriate 722*

Stem long or tall and sturdy; more than tri-seriate 723*

Stem short and thin or narrow; 1-3 seriate 724*

Stem long or tall and thin or narrow; 1-3 seriate 725*

Girder extends from bundle sheath to bulliform or colourless cells; not to epidermis 726*

Girder description continued:

Fibres interrupt the cells of the single or outer bundle sheath 727*

Fibres in contact with the cells of the single or outer bundle sheath 728*

CONTINUE ON FOLLOWING PAGE AND THEN RETURN AND DESCRIBE STRANDS AND GIRDERS OF ANY

OTHER ORDERS OF VASCULAR BUNDLES THAT ARE PRESENT IN THE SECTION.

R. P- ELLIS

83

ADAXIAL SCLERENCHYMA OF LEAF

726

727

726

84

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

7. SCLERENCHYMA OF LEAF CONTINUED

Continuation of circumvascular sclerenchyma surrounding, in contact with, or opposite the bundles; refers only to

longitudinal subepidermal sclerenchyma following the course above and below the vascular bundles. Complete description

is given of the circumvascular sclerenchyma associated with each of the different orders of vascular bundle present in the

leaf.

ABAXIAL sclerenchyma; sclerenchyma found below the vascular bundles:

No abaxial sclerenchyma present

Strand present; sclerenchymatous notin contact with the bundle sheath; separated by parenchyma or mesophyll

from sheath:

Minute strand consisting of only a few subepidermal fibres

Small strand; inconspicuous; epidermal cells unaltered under the strand:

Shallow, forming a strip; only 2-4 fibres deep; subepidermal

Narrow but fairly deep; only 2-4 fibres wide

As deep as wide as seen in T/S

Well-developed strand; conspicuous; epidermal cells under strand usually small and thick-walled:

As deep as wide as seen in T/S

Wider than deep as seen in T/S; in form of a band:

Arched, follows the shape of the abaxial rib

Straight, horizontal band

Girder present; sclerenchymatous in contact with, or interrupts the bundle sheath:

Very small girder; inconspicuous; epidermal cells usually unaltered under girder

Small girder; epidermal cells under the girder usually small and thick-walled:

Small, thin subepidermal strip

Narrow, deeper than wide

As deep as wide as seen in T/S; equidimensional

Well-developed girder; conspicuous; epidermal cells under the girder usually small and thickened:

Relatively wide and deep band; as wide or wider than the vascular bundle:

Arched, follow the shape of the abaxial rib

Straight, horizontal band

Narrowing towards bundle; triangular or trapezoidal

Relatively narrow and deep girder; narrower than the vascular bundle

Inverted T-shaped; horizontal band connected to bundle or bundle sheath by stem:

Stem short; shorter than horizontal cross-piece

Stem relatively long; as long or longer than horizontal cross-piece

Anchor-shaped; arched band follows shape of abaxial rib:

Stem short and sturdy; more than tri-seriate

Stem long or tall and sturdy; more than tri-seriate

Stem short and thin or narrow; 1-3 seriate

Stem long or tall and thin or narrow; 1-3 seriate

Continous or almost continuous abaxial hypodermal band

Girder extends from bundle sheath to bulliform or colourless cells; not to the epidermis

Girder description continued:

Fibres interrupt the cells of the single or outer bundle sheath

Fibres in contact with the cells of the single or outer bundle sheath

729*

730*

731*

732*

733*

734*

735*

736*

737*

738*

739*

740*

741*

742*

743*

745*

746*

747*

748*

749*

750*

751*

752*

753*

754*

N.B. CIRCUMVASCULAR SCLERENCHYMA OF EACH ORDER OF VASCULAR BUNDLE PRESENT IN SECTION

DESCRIBED SEPARATELY BEFORE COMMENCING WITH SCLERENCHYMA CELL STRUCTURE AND OTHER

SCLERENCHYMA TISSUE OF THE LEAF.

SCLERENCHYMA CELL STRUCTURE:

Fibres, circumvascular sclerenchyma comprised of thickened; elongated fibres:

Cell wall structure :

Very thick walled; lumen almost excluded

Cell walls of fibres of medium thickness; secondary wall distinct . . . .

Not markedly thickened; secondary wall not thick

Cell wall composition:

All cell walls lignified Le. stain red with saffranin

All cell walls of cellulose i.e. stain blue-green with fast green

Some sclerenchyma cell walls lignified others with cellulose walls:

In individual girder or strand groups

In different parts of the leaf section

Collenchyma; thickened parenchyma cells; wide lumens

755*

756*

757*

758

759

760

761

762*

86

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

7. SCLERENCHYMA OF LEAF CONTINUED

Continued from previous page and in contrast to the circumvascular sclerenchyma the types of sclerenchyma described

below are not described in relation to the different orders of vascular bundles.

SCLERENCHYMA BETWEEN BUNDLES:

Supernumary strands not associated with the vascular bundles but situated intermediate between them, usually beneath the

abaxial epidermis opposite the bulliform cell groups or furrows. Excludes sclerenchyma at the leaf margin and of the keel

or midrib.

DESCRIPTION:

No sclerenchyma present between the vascular bundles 763

Abaxial strands of sclerenchyma fibres present:

Opposite adaxial furrows with or without bulliform cells 764*

Opposite bulliform cell groups not associated with furrows 765*

Hypodermal band; sclerenchyma in form of more or less continuous abaxial band:

Extensions of girder or strand groups 766*

Hypodermal layer continuous 767*

SCLERENCHYMA IN LEAF MARGIN:

Includes all mechanical tissue situated in, or directly associated with the margin of the lamina. May incorporate or fuse

with the ultimate, the penultimate or other lateral bundles.

DESCRIPTION:

No sclerenchyma developed in association with the margin 768

Cap of sclerenchyma at the margin; not in contact with the lateral bundle:

Size of cap:

Very small; cpnsists of a couple fibres 769*

Relatively small; less than the width of a third order vascular bundle 770*

Well-developed; wider than third order vascular bundles 771*

Shape of cap:

Rounded cap 772*

Pointed cap 773*

Narrow, very pointed projection 774*

Crescent-shaped cap; sclerenchyma extends shortly along both abaxial and adaxial side of leaf 775*

Curved in shape with sclerenchyma extending along adaxial side of the leaf 776*

Curved in shape with sclerenchyma extending along abaxial side of the leaf 777*

Epidermal cells at margin:

Not fibrous; thin-walled or outer walls thickened; distinct from fibres 778

Small, fibrous; thickened on all walls; similar to fibres in T/S 779

Angular prickles of margin:

Enlarged bases present in margin 780

No prickle bases visible in margin 781

Nature of junction with mesophyll and remainder of lamina:

Adjoins normal mesophyll cells 782

Specialised enlarged parenchyma cells near cap or ultimate bundle 783

Lateral intercellular canal near margin; lysigenous duct not associated with the bundle 784

Hood of sclerenchyma; hood-like structure formed by sclerenchyma extensions above or below ultimate and other

lateral bundles; may or may not be in contact with bundles or bundle sheaths but must extend above or below or

beyond ultimate lateral vascular bundle:

Position of fibrous hood:

Fibres extend along adaxial side of the leaf 785*

Fibres extend along abaxial side of the leaf 786*

Extent of fibrous hood:

Short hood; extends as far as or over ultimate lateral bundle 787

Well-developed hood; extends as far as or beyond penultimate bundle 788

Penultimate bundle and associated sclerenchyma; may be no sclerenchyma development at the extreme margin but the

penultimate lateral bundle may be intimately associated with specialized girders, strands or bands:

Nature of lateral fibrous groups:

Bundle sheath of penultimate bundle modified and cells are horizontally elongated and in contact with the

normal sheath cells of the ultimate bundle 789*

Bundle sheath of penultimate bundle not modified but sclerenchyma continuous between the ultimate and

penultimate bundles from adaxial to abaxial epidermis 790*

Well-developed adaxial girder or band developed above the penultimate bundle 791*

Extensive adaxial band developed above the lateral 3 or 4 bundles 792*

SCLERENCHYMA ARRANGEMENT IN AC1CULAR LEAVES:

Refers to the sclerenchyma as seen in transverse section of permanently infolded leaves.

DESCRIPTION:

Abaxial strands present opposite the vascular bundles 793*

Abaxial girders present opposite the vascular bundles 794*

Continuous, abaxial, subepidermal band; not connected to the vascular bundles 795*

Continuous, abaxial, subepidermal band connected to bundles by girders 796*

Continuous, abaxial, subepidermal band and girders extending to adaxial surface 797*

Continuous, adaxial, subepidermal band not connected to the vascular bundles 798*

No, or very poor sclerenchyma development 799*

R. P. ELLIS

87

SCLERENCHYMA BETWEEN BUNDLES, AT MARGIN AND IN ACICULAR LEAVES

88

A

PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

THE POACEAE.

8. MESOPHYLL OF LEAF

Applies to all the ground tissue between the adaxial and abaxial epidermides. It usually consists mainly of the assimilatory

tissue or chlorenchyma but may be partly composed of translucent colourless cells. It does not include the bulliform cells

(which are part of the epidermis) the sclerenchyma or the bundle sheaths or their extensions.

CHLORENCHYMA

All the tissue containing chloroplasts excepting the parenchyma bundle sheath which may or may not contain chloroplasts

and is described under bundle sheaths. Does not include all the ground tissue between the adaxial and abaxial epidermides

especially if any translucent cells without chloroplasts are present.

DESCRIPTION:

Radiate chlorenchyma; radially arranged around the vascular bundles which are often close together; panicoid; may be

interrupted by sclerenchyma girders and thus divided into two groups on either side of the vascular bundle;

Number of layers of chlorenchyma cells arranged around the bundles:

Single layer of cells:

Tabular 801*

Isodiametric 802*

Two layers of radiating cells 803*

Numerous layers of radiating cells; cells long and narrow ; Isachne type 804*

Extent of radiating cells around bundles:

Radiating cells completely surrounding bundles 805*

Interrupted above bundles by sclerenchyma or colourless parenchyma girders 806*

Interrupted below bundles by sclerenchyma or colourless parenchyma girders 807*

Reduced to two strips of chlorenchyma by large girders or colourless parenchyma 808*

Relationships of successive radiating mesophyll groups to each other:

Separated by bulliform and colourless cell groups:

Completely divided; colourless cells continuous to abaxial epidermis 809*

Adaxially divided; colourless cells in adaxial half of leaf 810*

Separated by irregular chlorenchyma and intercellular air-spaces 811*

Radiating cells of successive bundles adjoin one another 812*

Indistinctly or incompletely radiate chlorenchyma; intermediate between radiate type and that not arranged in a

definite pattern:

Continuous between bundles 813*

Chlorenchyma divided by groups of colourless cells; tending to be radiate 814*

Irregular chlorenchyma; not radiate or arranged in a definite pattern ; vascular bundles usually widely spaced; festucoid:

Vertical arrangement in the mesophyll between successive vascular bundles:

Occupying the major or entire area between the adaxial and abaxial epidermides:

All cells of the chlorenchyma similar; homogenous:

Regular small cells; isodiametric; tightly packed 815*

Irregular; cells of different size and shape; often with intercellular air-spaces 816*

Cells pallisade-like in the adaxial chlorenchyma 817*

Chlorenchyma cells adjoining the bundles larger than those further removed 818*

Occupying the lower abaxial half of the leaf thickness 819

Occupying the lower abaxial third of the leaf thickness 820

Confined to small patches surrounding abaxial and adaxial grooves which contain stomata 821*

Horizontal arrangement in the mesophyll between successive vascular bundles:

Continuous between bundles:

Strap-shaped; horizontally elongated; bundles widely spaced 822*

Tall and narrow groups; bundles close together 823*

U-shaped; occupying the sides and bases of furrows or around bulliform cell groups 824*

Mesophyll groups divided by colourless cell groups 825*

Arm cells present or comprise chlorenchyma; walls invaginated:

Invaginations extend almost to opposite wall; cells divided into compartments 826*

Invaginations relatively short 827*

Fusoid cells present in mesophyll; translucent, elongated narrow collapsed cells; cavity between fusoid cells usually only

visible in T/S; bamboo type:

Successive fusoid cells separated by a vertical column of chlorenchyma one cell wide 828*

Successive fusoid cells separated by numerous chlorenchyma cells 829*

Distinctive cells, similar to parenchyma sheath cells but not associated with bundles present in mesophyll 830*

Lacunae or air spaces present in mesophyll:

Extent of lacunae :

Between all or most adjacent vascular bundles 831*

Continuous air spaces above third order bundles 832*

Aerenchyma associated with the lacunae:

Distinct lacunae bounded by compact mesophyll with no aerenchyma traversing the lacunae 833

Lacunae distinct but traversed by colourless aerenchyma cell chains 834

Irregularly defined air spaces in diffuse mesophyll with many intercellular spaces and chlorophyll-bearing

aerenchyma; often subtending the stomata 835

Stellate cells representing a diaphragm interspersed by sclerotic strands present 836

R. P. ELLIS

89

CHLORENCHYMA

>31

132

90

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

8. MESOPHYLL OF LEAF CONTINUED

COLOURLESS CELLS

Translucent cells constituting the remaining cells in the zones between successive vascular bundles, excluding the

chlorenchyma, the sclerenchyma and the bulliform cells. They vary in size and may be larger or smaller than the bulliform

cells but are always without chlorenchyma and never part of the epidermis i.e. not in contact with the surface.

DESCRIPTION:

Absent, no colourless cells present 837

Well defined groups of colourless cells present:

Cbsely associated with inflated bulliform cells or groups:

Size of colourless cells:

Smaller than bulliform cells; uninflated; often same size as chlorencyma cells 838*

Similar in size or shape to bulliform cells; inflated 839*

Width of extension of bulliform cell group; composed of colourless cells:

Narrower than bulliform cell group 840*

Same width as bulliform cell group 841*

Number of extensions from each bulliform cell group:

1 extension from each group 842*

2 extensions from each group; one on either side of the vascular bundle 843*

Extent of intrusion into the mesophyll by colourless cells:

Girders extend to the opposite epidermis 844*

Strands not extending to the opposite epidermis 845*

Band occupying the adaxial half of the leaf composed of colourless cells 846*

Not closely associated with large bulliform cells:

Origin at the base of furrows:

Girder-like extension to the opposite epidermis between the bundles 847*

Girder-like extensions to the bundle sheath or towards the bundle sheath 848*

Origin on flat epidermis or with slight furrows:

Girder-like extension extends to the opposite epidermis:

Column of girder only uni- or bi-seriate 849*

Girder multi-seriate 850*

Girder of colourless cells does not extend to the opposite epidermis 851*

III defined groups of smallish colourless cells; not associated with large or well defined bulliform groups; usually

adaxial:

Adaxial half of leaf consisting entirely of colourless cells 852*

Abaxial half of leaf consisting mainly of colourless cells 853*

Present on adaxial side of small vascular bundles only 854*

Form arches over third order bundles and extend to the abaxial epidermis 855*

Abaxial irregular groups; may be associated with sclerenchyma strands 856*

DESCRIPTION OF INDIVIDUAL CELLS:

Structure of colourless cells:

Inflated, large parenchyma cells; resemble bulliform cells in size and shape 857

Smaller, thin-walled parenchyma cells; rounded in shape:

Cells regular in size and shape ; tissue uniform in appearance 858

Cells irregular in size and shape; tissue irregular in appearance 859

Thick-walled parenchyma or collenchyma cells 860

R. P. ELLIS

91

COLOURLESS PARENCHYMA CELLS

92

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

9. EPIDERMAL CELLS IN TRANSVERSE SECTION

Includes all cells and their appendages present in the epidermal layer. Can be subdivided into the bulliform cells and typical

thickened epidermal cells.

ADAXIAL AND ABAXIAL EPIDERMIS of the leaf; cells of the upper and lower epidermis considered separately and the

adaxial epidermis is fully described before commencing with the abaxial epidermis:

Cells of adaxial epidermis 901

Cells of abaxial epidermis 902

BULLIFORM CELLS

Colourless cells forming part of the epidermis but differing from the remaining epidermal cells by being markedly larger

and inflated. Characteristically they are large and are restricted to groups often intimately associated with colourless cells

but all gradations are found. Thus small solitary bulliform cells may be present or the epidermis may be comprised

primarily of large, rounded and inflated cells. All these variations are included under bulliform cells for descriptive

purposes. For bulliform cells of the keel see under midrib and keel.

DESCRIPTION: Adaxial and abaxial epidermides considered separately.

Absent, no bulliform cells in epidermis being described; consists of small cells; often thickened 903*

Bulliform cells only; not arranged in regular groups; epidermal cells rounded and inflated the same size or slightly larger

than the cells of the bundle sheath:

All or most epidermal cells of this type 904*

Isolated cells larger; may comprise irregular small groups:

On sides of deep furrows 905*

Not in furrows; usually in leaves with flat surfaces 906*

Groups of bulliform cells present in epidermis; well-defined and regular; distinct from normal epidermal cells:

Small bulliform cells; not conspicuously larger than the normal epidermal cells:

Not associated with colourless cells 907*

Closely associated with colourless cells; indistinct 908*

Extensive groups of large, inflated bulliform cells extending over one or many vascular bundles:

Not associated with colourless parenchyma:

Distribution of extensive bulliform groups:

Present throughout the epidermis; may be slightly reduced opposite the larger bundles 909*

Present in most of the epidermis but not present opposite the first order bundles and usually reduced over

the second order bundles 910*

Size of constituent cells; taken in areas opposite the third order bundles:

Occupy less than 5 of the leaf thickness 911*

Occupy more than £ of the leaf thickness 912*

Associated with colourless parenchyma cells together with which it forms a zone of colourless cells above or

below the chlorenchyma; may be disrupted over the first order bundles:

Colourless zone comprises more than \ of the leaf thickness 913*

Colourless zone comprises less than ^ of the leaf thickness 914*

Restricted groups of large, inflated bulliform cells; Zea type; parallel-sided cells i.e. inner tangential wall same length

or only slightly shorter than the outer tangential wall:

Projecting above the level of the epidermis; cf. cushion cells of macro hairs 915*

Level with the general epidermal surface 916*

Restricted groups of tall and narrow bulliform cells; not inflated; parallel-sided cells or with outer tangential wall

slightly shorter than the inner wall; lateral walls long and straight 917*

Fan-shaped groups; each cell of group with outer tangential wall shorter than inner tangential wall; median cell of

group appreciably larger than the remainder:

Situated at bases of furrows 918*

Not found at bases of furrows or present in leaves without furrows; some cells parallel-sided 919*

Resemble fan-shaped groups due to central cell of group being larger and of a different shape than the rest which

may be small by comparison; not necessarily with shorter inner tangential walls; Sporobolus type:

Shape of central cell of group which is always narrow at epidermis and often recurved or straight at area of

contact with lateral bulliform cells of group:

Inflated, rounded; short area of contact with lateral cells 920*

Inflated, fan-shaped 921*

Narrower than deep; shield-shaped 922*

Elongated with parallel sides 923*

Elongated rather pointed base; diamond-shaped 924*

Size of central cell:

Relatively small; not much larger than bundle sheath parenchyma 925*

Occupy 5 - 5 of the leaf thickness 926*

Occupy more than 5 of the leaf thickness 927*

Narrow groups of bulliform cells and intimately associated large colourless parenchyma cells penetrating deep into

the mesophyll; the contact with the adaxial epidermis made by the bulliform cell unclear; Arundo type 928*

Bulliform cells and closely associated colourless cells forming an extensive column or girder extending from the base

of an adaxial furrow deep into leaf:

Nature of column or girder:

Column uni-seriate 929*

Column bi-seriate 930*

Column multi-seriate 931*

Position of column:

Extends from base of furrow to abaxial epidermis 932v

Extends from base of furrow towards a vascular bundle 933”

CONTINUE DESCRIPTION OF ADAXIAL EPIDERMIS OVERLEAF AND THEN RETURN AND DESCRIBE THE

ABAXIAL EPIDERMIS.

R. P. ELLIS

93

BULLIFORM CELLS

CCPTOP

rcmJILrnir

907

otoh wnn ™ amn

©@@U>@@ 3 © @0

913

914

917

925

929

927

94

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

9. EPIDERMAL CELLS IN TRANSVERSE SECTION CONTINUED

TYPICAL EPIDERMAL CELLS AND APPENDAGES

Includes all epidermal structures as seen in transverse section excluding the bulliform cells. Incorporates the cuticle,

macro-hairs, prickles and papillae as seen in transverse section.

DESCRIPTION:

Cuticle and Thickening of epidermal cell walls:

Outer tangential wall:

Outer walls not thickened and with a very thin cuticle

Outer walls slightly thickened or with a thin cuticle

Outer walls thickened and covered by a distinct, thick cuticle continuous over the epidermal cells:

Cuticle and cell wall equal to or greater than the depth of the average epidermal cells

Cuticle and cell wall less than the depth of the average epidermal cells

Outer tangential wall of each epidermal cell thickened individually:

Outer wall occupies half or more of the depth of the cells

Outer wall occupies less than half of the depth of the cells

Irregular; cells of different sizes comprise the epidermis

Radial and Inner tangential walls of epidermal cells:

All walls of epidermal cells thickened

Only outer wall thickened

No walls thickened

Macro-hairs as seen in transverse section; nature of hair bases described:

No macro-hairs visible

Superficial bases; not sunken into the leaf or embedded between inflated bulliform cells:

Epidermal cells not modified to form cushion cells associated with the base of the hair:

Swollen base not much larger than and situated between normal epidermal cells; included here are probably

some elongated prickle hairs:

Hair short and thick

Hair slender and elongated

Constriction above bulbous base embedded in normal epidermal cells:

Hair very thin and slender

Hair thickened and stiff

Epidermal and other cells modified to form a cushion base:

Base surrounded by small thickened cells which may extend a short way up the hair:

Hair with enormous swollen base

Base of hair not exceptionally large and swollen

Sunken bases; sunken in leaf or embedded between large, inflated epidermal cells or bulliform cells:

Constriction above bulbous base embedded between large epidermal cells:

Size and shape of hairs:

Hairs slender but relatively short

Hairs thick and short

Hairs long and very slender

Hairs long and relatively thick

Nature of bulliform or epidermal cells surrounding base:

Raised around hair forming a definite cushion

Unraised bulliform cell groups

Base bulbous and not constricted; embedded between bulliform cells

Prickles and Hooks as seen in transverse section:

No hooks or prickles visible

Small hooks present in the epidermis; usually located in intercostal areas:

With short, curved barbs

With straight, pointed and slender barbs:

Present between typical epidermal cells

Present between inflated bulliform-like cells

Thickened prickles present in epidermis; usually located opposite the vascular bundles:

Pointed broad prickle; base not bulbous

Small with short barb

Bulbousbase; barbed

Bulbous base; no barb; asperite

Papillae on the epidermal cells as seen in transverse section:

Shape of outer tangential wall of the epidermal cells:

Outer walls of epidermal cells not arched to any marked degree; flattened

Outer walls of epidermal cells arched but not papillose

Arching of outer wall exaggerated; papillose; inflated papillae as wide as epidermal cells:

Thin-walled wide papillae scattered throughout the epidermis

Entire or major part of epidermis composed of thin-walled wide papillae

Papillae wide; as wide as or slightly narrower than epidermal cells; not inflated and thin-walled:

Distal, outer wall markedly thickened

Papillus thickened and cuticular

Papillae narrower than the epidermal cells:

Size of papillae:

Relatively broad but not much more than half the width of the epidermal cells

Much less than half the width of the cell

Number per cell as seen in transverse section:

One per cell

Bifurcate; two per cell

N.B. ADAXIAL AND ABAXIAL EPJDERMIDES DESCRIBED SEPARATELY.

934*

935*

936*

937*

938*

939*

940*

941

942

943

944

945*

946*

947*

948*

949*

950*

951*

952*

953*

954*

955*

956*

957*

958

959*

960*

961*

962*

963*

964*

965*

966*

967*

968*

969*

970*

971*

972*

973*

974*

975*

R. P. ELLIS

95

EPIDERMAL CELLS

ecu mi rm nm mt inr y&to

mm CE «E»Cte flVr tCt tat

966 968 970 972 974

mm tnnxi AdMbtOcc Mtl mm

967 969 971 973 975

96

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

ANATOMICAL CHARACTERS

The following presents a brief conspectus of the

nature of the various structures used in describing

grass leaf anatomy, as well as their range of structural

variation. Where relevant, an account is given of

the function or development of these structures and

the factors which contribute to their variability.

Their importance as either taxonomic or diagnostic

characters is stressed.

Outline of the lamina in transverse section

The foliage of the grass plant is comprised of the

sheath, the ligule and the leaf-blade. In the following

account, the leaf-blade, the expanded and free

portion of the leaf as distinct from the sheath, is

termed the lamina or the blade.

Outline of the lamina of open leaves

In many grasses with open leaf-blades the two

halves of the lamina on either side of the median

vascular bundle, midrib or keel, are relatively thin

and wide, of equal width and usually symmetrically

arranged about the median region. Infrequently,

however, in certain species the portion of the lamina

to one side of the midrib may not be symmetrical

with that on the other half or perhaps the leaves

may be relatively narrow and rather thick. These are

exceptions to the general rule but are nevertheless

useful for indentification purposes.

The lamina of most grass leaves in transverse

section appears flattened and expanded and a straight

line connects both margins and the median bundle

or keel. However, expanded laminae may become

variously altered by involution usually by becoming

inrolled or infolded. Infolding results in the

appearance becoming V- or U-shaped because the

two lateral halves of the lamina tend to become

folded adaxially towards each other on either side

of the median vascular bundle which may, or may not,

be adaxially grooved or incorporated into a keel.

Grooves or keels serve to accentuate the V- or

U-shape which characterizes many species. Inrolled

leaves may appear variously U-shaped or may show

either convolute inrolling from one margin only, or

involute inrolling where the leaf-blade rolls inwards

from both margins.

The degree of infolding or inrolling varies with the

environmental conditions and thus is not of much

value diagnostically (Metcalfe, 1960). However, leaves

of grasses that are fully expanded under optimum

conditions invariably exhibit a characteristic type of

involution in response to environmental stress.

Therefore, it is the nature, and not the degree, of the

infolding or inrolling that is of importance

taxonomically and diagnostically. Furthermore, it

must be noted that some leaves remain flat on drying

and in others the leaf folds inwards a little near the

keel but the margins remain reflexed. Other responses

are shown by the corrugations becoming more pro-

nounced in species with plicate blades, and the pro-

duction of filiform leaves in species which normally

possess expanded blades. Further details of the

involution process are included under bulliform and

colourless cell function.

Expanded leaves of different species thus exhibit

various, but constant, movements in response to

adverse environmental change, the resultant shape

being of specific or generic diagnostic value. Various

members of the Andropogoneae (Hyparrhenia, Cym-

bopogon and Dichanthium) often show limited

infolding near the keel with the margins remaining

reflexed and several species of Cymbopogon produce

both expanded and filiform leaves (Vickery, 1935).

Some Brachiaria and Opiismenus species do not alter

shape at all under conditions of water stress and

remain flat and expanded. Certain grasses even show

nyctinastic movements as described for Leersia

hexandra by Goebel (Bor, 1960) in which the leaves

are folded in dull weather but flatten in bright sunlight.

The variability exhibited by the leaf outline

demonstrates the necessity for a sampling method

which will adequately reveal the extremes of this

variation. This should be achieved if adequate

numbers of specimens of each species are collected,

and fixed immediately, from different areas, over an

extended period of time and from as many habitats as

possible.

It appears reasonable to postulate that the type of

folding adopted by the mature leaf in response to

environmental conditions, as well as the symmetry of

the leaf, may be correlated to the way in which the

leaf is folded in the bud. Thus, Amedei (1932) has

shown that Chaetochloa palmifolia ( =Panicum palmi-

folium) is folded conduplicately in the bud and the

mature leaf is more or less pleated depending on

circumstances. The blades of grass leaves may be

either folded, convolute or pleated in the bud (Bor,

1960). Therefore, the bud arrangements may simply

be linked to the major types of involution exhibited

in mature leaves except, perhaps, involute inrolling

which is difficult to explain. Does it happen then, that

a leaf which is convolute in the bud will exhibit

definite V-shaped movements as a mature leaf

instead of the expected convoluted inrolling? This

possibility merits further investigation.

Apart from the characters of the shape of the

outline, under optimum and adverse conditions,

quantitative data of leaf width and thickness are also

of value in describing and identifying leaves of

different grasses. De Winter (1951) mentions the

extreme thinness (about 0,05 mm) of the leaf of

Prosphytochloa prehensiiis (= Potamophila prehensiiis)

where the leaf is only about four cell layers thick. These

quantitative characters are subject to considerable

variation, and categories must be broadly defined to

accommodate this variability. Detailed measurements

have not been included in the keys and measuring

is done by estimation of the leaf dimensions in

relation to the known diameter of the field of view

of the microscope used in the study. These broad

categories are, nevertheless, useful for comparison

with measurements given by other authors and can

be easily modified as required.

Outline of permanently infolded leaves

In many grasses the lamina is very strongly and

often permanently infolded or involuted. In such

leaves the stomatal bands are invariably restricted

to certain areas of the adaxial surface, usually on the

sides of adaxial furrows within the channel. A thick

cuticle and extensive sclerenchyma development is \

characteristic although this is not universally true as

shown by Lewton-Brain (1904) for Mibora minima

( =Mibora verna) which is entirely parenchymatous, i

However, these leaves can be distinguished by the

fact that they are extremely narrow and permanently

infolded to the extent that the internal structure

is altered.

Numerous leaf-types defined on morphological

criteria fall into this group. Examples are setaceous

(wiry and bristle-like), filiform (thread-like), acicular

(needle-like), junciform and cylindrical leaves.

Anatomically these different types cannot be separated

with any degree of certainty and a term inclusive of

all permanently infolded leaves is required.

R. P. ELLIS

97

Acicular leaves invariably fulfill the requirements

of the definition of being permanently infolded to the

extent that the internal structure is altered. In filiform

leaves, often produced under adverse conditions by

species which normally have open, expanded leaves,

however, the blade is reduced to an enlarged midrib

structure, with a mass of colourless parenchyma above

the bundles and the lateral laminae being reduced or

absent. Some species of Cymbopogon exhibit this

reduction regularly. The extreme case of this

specialization is seen in Miscanthidium teretifolium

I which is cylindrical in transverse section and the

adaxial surface can be recognised only by the presence

of a minute groove on the adaxial side of the cylinder

(Metcalfe, 1960). In these cases, it must be stressed,

the internal structural alteration of the leaf does not

occur as a result of the leaf being permanently folded

but rather as a result of loss of the lateral portions

of the blade and a corresponding development of

the midrib.

The cylindrical condition can probably arise by

one of two different processes: the loss of the lateral

regions and a concurrent development of the keel or

by parenchyma development in a permanently infolded

leaf excluding the adaxial channel. For convenience

these two types of narrow leaf are included together

under permanently infolded leaves. This further

illustrates the need for a single term describing all

reduced, permanently infolded and/or structurally

altered grass leaves.

Transverse sections of these leaves often exhibit

characters that are of taxonomic value because in

assuming the permanently infolded, acicular form,

the leaves of grasses of different affinities have not

acquired precisely the same arrangement of vascular

and mechanical tissue (Metcalfe, 1960). The outline

and the shape of the adaxial channel and ribs and

furrows may be distinctive as well. Burbidge (1946a)

was able to separate ten species of Triodia, all

possessing acicular leaves, using characters of the

outline, the adaxial channel and adaxial and abaxial

furrows. For descriptive purposes it is, therefore,

necessary to describe the shape of the blade as

outlined by the abaxial surface, to count the number

of vascular bundles present in the section, to describe

the adaxial channel and to include leaf width and

thickness dimensions. Ribs and furrows and

sclerenchyma are described in later “keys”. With

these descriptive criteria the differences present in

permanently infolded leaves are of significant specific

diagnostic value.

It is absolutely essential to standardize on the

portion of the blade to be sectioned in permanently

infolded leaves as the shape can vary from the base

to higher up the leaf. Arber (1923, 1934) noticed

that folded leaves may approach radial symmetry

and become cylindrical in the apical regions. In

Triodia (Burbidge, 1946a), towards the apex of the

lamina the lateral veins disappear one by one,

commencing with the marginal pair. There is a

corresponding progressive reduction in the amount

of mesophyll so that finally the pungent point is

formed of the median vascular bundle surrounded

by sclerenchyma. This progressive change, of many

characters, along a single blade is equally important

in open, tapering leaves and is the prime reason for

standardizing on material from the central region of

the lamina from ligule to tip.

In the cases of some narrow, folded leaves it is often

debatable whether they in fact should be described

under open or under permanently infolded leaves.

In such cases it is perhaps safest to describe them

under both of these arbitrary categories.

Longitudinal ribs and furrows

The adaxial and abaxial surfaces of the grass leaf

blade may be either flat or longitudinally ribbed.

The ribs are usually developed in association with and

adjacent to the larger vascular bundles. They are

generally characteristic of and more fully developed

on the adaxial than the abaxial surface (Metcalfe,

1960). Ribs have also been termed ridges (Arber,

1934; Wilson, 1971) and if ribs are present there

must be corresponding furrows between adjacent

ribs. Furrow, in the context used here, is a term

reserved exclusively for the depressed intercostal zone

between adjacent ribs and which, with few exceptions,

occur between the vascular bundles. If furrows are

present in any other situation they are termed grooves

to avoid possible confusion. Examples are the grooves

present at the apex of the well-developed ribs of

Sporobolus artus (Goossens, 1938) or those described

immediately abaxial to the larger veins described

by Sabnis (1921). In permanently infolded leaves the

area bounded by the infolded lamina is characterized

by being referred to as a channel.

In different species ribs and furrows may vary in

height or depth, transverse shape, spacing and

location. The ribs of a single grass blade can be of

one or more distinct sizes or shapes, the ribs of each

type being consistently associated with a specific order

of vascular bundle. It is possible to measure the

inclination of the ribs by measuring the angle formed

by the sides of the two adjacent ribs at their base in

the furrow (Wilson, 1971). Consequently, leaves with

obtuse angles would have flatter surfaces than those

with acute angles. The abaxial ribs may sometimes be

taller than those of the adaxial surface. Where well-

developed abaxial and adaxial ribs and furrows are

similar and occur opposite each other the leaf section

resembles a string of beads and is termed moniliform.

There are numerous examples of species of grasses

possessing ribs or furrows that do not conform to the

generalized definition outlined above. Lewton-Brain

(1904) illustrates the enormous triangular ribs of

Aira caespitosa where three vascular bundles are

included in a single rib. Glyceria fluitans, another

example illustrated by Lewton-Brain (1904), possesses

low adaxial ribs situated between the vascular bundles

which are situated beneath the ‘furrows’. The ribs

are occupied by copious air spaces. The flat-topped

abaxial ribs of Digitaria macroglossa are exceptional

because the ribs are in fact sclerenchyma caps and

are not linked to the vascular bundles in any way.

Metcalfe (1960) comments on the correlation

between prominent ribs and a marked capacity for

involution, and states that this ability has been evolved

in response to ecological conditions. Furthermore, this

has occurred independently in various grasses and con-

sequently leaves of this type are found in grasses

between which there are no close taxonomic affinities.

Rib development is nevertheless, often useful for

specific diagnostic purposes and Gordon-Gray and

Ward (1970) have contrasted the consistent flatness

of the leaf surfaces of Phragmites mauritianus with

the slight ribs developed on comparable surfaces of

leaves of the closely related P. australis.

A paper by Wilson (1971) shows that the adaxial

ribs of Lolium perenne appear to be under the control

of a few genes as indicated by the rapid and immediate

response to selection achieved in this study. Thus,

in a species which appears to exhibit a wide range of

phenotypic variation with respect to rib development,

this variation may in actual fact be the direct mani-

festation of the genotype. This fact should be borne

in mind when variation patterns of adaxial ribs are

studied.

98

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

Median vascular bundles, midribs and keels

In order to eliminate any possibility of ambiguity

arising from the use of these terms it is necessary to

define them in the context that they are to be used

here. The median vascular bundle is so termed only

if the centrally situated vascular bundle of the leaf

is structurally indistinguishable from the other basic

or first order vascular bundles of the leaf, and if this

median bundle is not associated with any parenchyma

development or thickening of the leaf. On the other

hand, a midrib is centrally positioned but structurally

distinct from the other first order bundles with respect

to size, bundle sheath, sclerenchyma girder or vascular

structure and lacks associated parenchyma. When

parenchyma or bulliform cells are developed in

association with the median bundle or bundles, the

whole structure, often incorporating many bundles,

is termed a keel. The median bundle of a keeled leaf

may be a midrib, and thus distinguishable from first

order bundles, or a median vascular bundle, being

indistinguishable from the other first order vascular

bundles. A keel usually projects either abaxially,

adaxially or both, but this is not necessarily so and

in V-shaped leaves, unless there is a marked thickening

in relation to the rest of the lamina, the median bundle

alone is considered to comprise the keel, if it is

accompanied by parenchyma.

This distinction of the three distinct types is con-

sidered necessary because in the past these terms have

been interchanged by various workers. Metcalfe (1960)

considers a projecting midrib, whether associated with

parenchyma or not, to constitute a keel. Thus, no

distinction is made between leaves with or without

parenchyma in association with the median vascular

bundle.

The size of the keel in an individual leaf varies,

usually becoming progressively larger towards the base

of the blade so that, in transverse section, the extent

to which a keel is prominent depends on the level at

which the section is taken. Therefore, the central

part of the leaf must be used for comparative purposes.

Vickery (1935) states that, in addition to variation

at different points along a single leaf, the size and

degree of development of the keel varies tremendously

in different leaves of the same species, so that the

appearance in transverse section can only be of

importance when taken in conjunction with the

macroscopic appearance of the keels of a number of

leaves. This is definitely not necessary in some groups

and Fisher (1939) found the keel-size, amount of

parenchyma and sclerenchyma and the bulliform

cells — to be a valuable diagnostic character in

separating four species of Chloris.

Keel structure, although subject to variation, is

important diagnostically because the various types of

keel that are recognised are easily distinguished

except that the inconspicuous keel with one vascular

bundle merges into that with three bundles. Thus,

size may vary but shapes are constant and distin-

guisable. In addition there are numerous associated

characters such as sclerenchyma, parenchyma and

bulliform cells of the keel, air spaces and the nature of

the adaxial side of the keel.

Bambusoid and oryzoid grasses display distinct keel

structure with prominent lacunae, with or without

diaphragms (Holm, 1896), and with a complex system

of bundle arrangement. The keel is comprised of two

or more vascular bundles distributed near both the

abaxial and adaxial surfaces and sometimes in the

interior as well. This characteristic structure has been

discussed by various authors such as Holm (1892,

1895, 1896), Arber (1934), Jacques-Felix (1955),

Schweickerdt & Marais (1956), Tateoka (1965) and

Launert (1965). In addition to the Bambusae &

Oryzeae, the Streptochaetae and Prosphytochloa

prehensilis (= Potamophila prehensilis ) in South Africa

(de Winter, 1951) possess superposed adaxial bundles

in the keel. Air spaces developing in the keels of older

leaves are common in many other genera. Sporobolus

artus var. lysigenatus and S. pyramidalis (Goossens,

1938) are examples.

Vascular bundle arrangement

The vascular bundles, or parallel longitudinal veins,

are usually arranged in a single row embedded at

various positions in the mesophyll as seen in section.

Certain grasses, such as Oryza coarctata (Tateoka,

1963) and Porteresia (Tateoka, 1965a) have a normal

abaxial vascular bundle in each rib, but in addition, a

superposed, adaxial, amphivasal bundle is situated

immediately anterior to each of the abaxial bundles.

This type of arrangement is also found in the keels

of some bambusoid and oryzoid grasses.

The arrangement most commonly found in grasses*

is for all the bundles to be centrally positioned in the

vertical plane of the blade. However, all bundles may

be located closer to the abaxial or to the adaxial

surface. Irregular arrangements, with bundles of

different sizes or orders situated at different levels

within the mesophyll are also found, In these instances,

a regular pattern of arrangement of the different i

orders of bundles in the mesophyll is usually dis-

cernible. Cases where the bundles are located at

irregular and inconsistent levels are rare, such as |

Stipa tenacissima (Metcalfe, 1960). This positioning j

of the vascular bundles in the blade appears to be a j

useful diagnostic character above the genus level that

has been largely overlooked in the past.

The total number of vascular bundles in a section

through a leaf blade taken halfway between the

sheath and apex is inconsistent, varying with the

width of the leaves. Little reliance can be placed on

the total number of vascular bundles in a leaf as

Vickery (1935) correctly stresses. She found that

Themeda avinacea and Cymbopogon exaltatus may

have flat expanded leaves 5-7 mm wide but under

certain conditions filiform leaves consisting essentially

of an enlarged keel are formed. This reduction in the

width of the blade involves a reduction in the number

of vascular bundles. Wider leaves do not necessarily

have more vascular bundles than narrower leaves,

however, as the bundles may be more or less crowded

in different specimens of the same species. The total ;

number of vascular bundles in a section is only

considered to be useful in permanently infolded leaves,

where there has been a reduction in the total number

of vascular bundles.

The relative proportion and the alternation of the

various sizes of vascular bundle along leaves of any

one species of grass is remarkably constant. Slight

variations in this respect may be found in leaves

taken from different levels on a single plant, the

variation being linked with differences in the width

of the blades. For this reason basal leaves have been

standardized upon for comparison. However, the

proportion of first order bundles and smaller bundles

in grasses belonging to different genera and species

show marked differences.

The distribution patterns of bundles of different

orders, and their variation, are important diagnostic

characters. Thus, Holm (1891a) found a pattern of

1 first order bundle, 1 third order bundle, 1 second

order bundle, 1 third order bundle, 1 first order

R. P. ELLIS

99

bundle, etc., in Uniola latifolia. He notes that this

formula is not strictly constant but that it gives the

general features concerning the relative number and

arrangement of bundles. For other species of Uniola

he was unable to give a formula because of excessive

variation, especially in leaves from different localities.

Leigh (1960, 1961), in a study of a few Eragrostis

curvula strains distinguished two groups both with a

basic pattern of 3 or 4 second order bundles between

successive first order bundles repeating itself either

four or eight times along the width of the leaf.

The use of a formula to describe the pattern of

arrangement of the various orders of vascular bundle is

only suitable when a regular pattern from median

bundle to margin is present. Often no pattern is

obvious with progressively fewer first order vascular

bundles and more third order vascular bundles nearer

the margin or more first order vascular bundles and

fewer second and third order vascular bundles

laterally situated. For this reason the use of formulae

to describe these patterns is not suitable. On the

other hand, if a constant part of the blade is described,

e.g. the central part of the lamina between the median

bundle and the margin, these observations on bundle

alternation will be comparable, even for leaves where

the pattern changes from median bundle to margin.

Vascular bundle structure

Metcalfe (1960) visualizes the vascular bundle as

consisting solely of the xylem and phloem elements,

the bundle sheaths not being treated, for descriptive

purposes, as if they constitute part of the bundle.

For consistency and convenience this principle has

been adopted in the present work and thus, when a

bundle is described as being circular or angular in

outline, these terms refer only to the outline of the

xylem and phloem tissue.

The vascular bundles of the Poaceae are divided

into three orders or ranks for descriptive purposes.

These orders are not necessarily distinct but may

intergrade and intermediate types are sometimes

present. However, within a given leaf, three, and

sometimes even four, classes of bundle are usually

evident even although their structure may not con-

form exactly with that of the generalized definitions

given below.

The three different orders in a single leaf are usually

immediately evident because they differ in size.

However, the relative diameters of the various orders

cannot be used satisfactorily for the grass family as

a whole, because equivalent orders of bundle differ in

size and structure from one species to the next.

Bowden (1964) classified bundles using the diameter

as a criterion, but in species where all bundles are

approximately the same size, but structurally different,

this system is inadequate.

The first order bundles, or basic type (Metcalfe,

1960) are characterized by having a metaxylem vessel

on either side of the protoxylem which may be non-

functioning, and replaced by a lysigenous cavity or

lacuna. A few well-developed protoxylem vessels may

be present in addition to the lysigenous cavity, or

both cavity and vessels may be absent. In the second

order bundles the xylem and phloem are easily

distinguishable, but the large metaxylem vessels are

lacking as is the lysigenous cavity. The third order

vascular bundles are often small bundles in which the

xylem and phloem is reduced to a few elements or

may even be indistinguishable using light microscopy.

In first order bundles the metaxylem vessels may

appear to be paired, but these are merely the oblique

overlapping ends of otherwise solitary vessel elements.

Thus the number of metaxylem vessels in an

individual bundle is unreliable as a diagnostic

character because it varies with the level at which the

sections are taken (Metcalfe, 1960). For this reason,

only the size and shape of these vessels are mentioned

in the descriptive keys.

The phloem of the first order vascular bundles

may exhibit varying degrees of sclerosis. This is

especially common in grasses from dry localities

(Metcalfe, 1960). In Triodia (Burbidge, 1946a) and

Merxmuellera {— Danthonia ) (de Wet, I960) the

phloem may be divided into two or three groups by

intrusion of small fibres. The phloem may adjoin the

inner or parenchyma, bundle sheath or it may be

completely surrounded by thick-walled fibres which

jsolate it from the xylem and the bundle sheath

(Jefferies, 1916).

Bundles of the second order commonly resemble

the first order bundles in shape and size and are only

recognisable by the absence of the metaxylem vessels.

Some authors have pointed out that second order

bundles may show a tendency to develop into first

order bundles (Vickery, 1935; Goossens, 1938).

In third order vascular bundles the metaxylem

vessels are always lacking and in the smallest

bundles of this order the xylem and phloem elements

may be indistinguishable. These bundles, in fact,

form a reduced conducting system and sometimes

only a few lignified cells are present together with the

phloem elements. Fisher (1939) notes that in these

instances neither the protoxylem nor the metaxylem

is present. In some genera the third order bundles

are not conspicuously smaller than the basic type

bundles, but are then distinguishable by the absence

of sclerenchyma girders, or even strands, and/or the

presence of bulliform cells adaxially.

The shape of the third order bundles may be

classified as angular, inconspicuously angular, circular

or elliptical in outline. They exhibit these shapes more

clearly than do the larger bundles. This shape is

taxonomically an important character, linked with

the size and number of bundle sheath cells. Thus,

small, angular bundles are, on the whole, characteristic

of the panicoid, and non-angular bundles of the

festucoid grasses.

Various classifications of the orders of vascular

bundles employ the extent of development of the

sclerenchyma strands and girders. Thus, Breakwell

(1914, 1915) distinguishes primary bundles, in which

the sclerenchyma is in direct contact with the bundle,

and secondary bundles which possess an entire bundle

sheath. Bowden (1964) also groups first and second

order bundles as those possessing sclerenchyma girders

and third and fourth order bundles as those with

strands. As was pointed out above, the nature of the

sclerenchyma developed in association with the

various orders of vascular bundle is especially

important in recognizing the larger types of third

order bundle, but as bundles are defined here, bundle

sheaths and sclerome are not included in vascular

bundle descriptions.

Surprisingly few detailed studies have been con-

ducted on the phloem of the Poaceae and these only

recently. Thus, the mature phloem of A vena was

studied by O’Brien & Thinman (1967), Buvat (1968)

studied the sieve elements of Hordeum and Singh &

Srivastava (1971) the phloem of Zea.

These studies have shown that the first-formed

protophloem elements are located abaxially within a

bundle and subsequent sieve elements differentiate

successively in an adaxial direction to within a

couple of cells of the differentiating xylem elements

Thus, in a mature bundle, all the major features of

100

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

differentiation of the sieve elements and companion

cells are visible. The sieve elements of the protophloem

often lack companion cells and may differentiate

without lateral expansion. The later-formed sieve

elements, as a rule, are accompanied by one or more

companion cells and extent laterally as a first step in

differentiation. Furthermore, the protophloem is

usually thin-walled and may become crushed in

mature bundles. The metaphloem has thicker walls

and may even be thick-walled as in Distichlis stricta

(Roy, 1969).

The xylem consists of primary xylem only. In first

order bundles it can be divided into the protoxylem,

which is that area in which the tracheary elements

have either annular or helical thickenings or both,

and metaxylem, which is the remaining primary

xylem. This metaxylem can be sub-divided into the

early and the late metaxylem; the late metaxylem

being the two large vessels so characteristic of these

bundles.

The structure of these late metaxylem vessels is an

indication of their specialization. Cheadle (1955,

1960) has shown that specialization in the Angio-

sperms proceeds from long vessel members, that are

angular as seen in transverse section, and with long

scalariform plates with many bars and perforations

on oblique end walls, to short vessel members that are

circular in section, with simple plates, with single

perforations on transverse end walls. In the Poaceae

no great variation occurs in the late metaxylem which

is very specialized. Thus, simple plates are common

and scalariform plates rare. A character that is

variable, and revealed in section, is whether or not the

vessel members are circular or angular.

Vascular bundle sheaths

In the Poaceae every vascular bundle is surrounded,

either completely or partially, by one or two single

layered bundle sheaths. These bundle sheaths are

taxonomically very important, and as a general rule

single sheaths are characteristic of the panicoid and

double sheaths of the festucoid grasses. Structural

variations of the bundle sheath are also important for

diagnostic purposes, as well as constituting evidence

for important physiological differences between the

various groups of tribes of the Poaceae. As would be

anticipated in a tissue undergoing extensive changes,

even within genera, to almost every generalization

there are exceptions. Notwithstanding, the bundle

sheath is undoubtedly of the utmost importance from

a comparative anatomical viewpoint and fortunately

numerous ultrastructural, physiological and taxo-

nomic studies of the structure and functions of the

sheath have been undertaken.

Recent terminology, to avoid reference to physiolo-

gical activities and anatomical structure, as well as to

eliminate the concept of specific functions for the

two layers (Lommasson, 1957), has been restricted to

the terms double (comprised of an inner and outer)

and single sheaths. Various other terms have been

used in the past, but none of them are applicable to

the grass family as a whole.

The single sheath, and usually the outer sheath,

when there are two, commonly consists of a single

layer of large, thin-walled or slightly thickened cells.

In section they are generally inflated and conspicuous,

being larger than the adjoining mesophyll cells.

Panicoid grasses, in general, have more inflated single

or outer sheath cells than those of the outer sheath

of the festucoid species (Metcalfe, 1960). This sheath

has often been referred to as the parenchyma bundle

sheath but other terms include starch sheath, border

parenchyma as well as mestome sheath. None of

these terms are generally applicable, as for example

Aristida species have an inner bundle sheath con-

sisting of larger cells than the outer sheath, which

may be composed of thickened cells.

The typical single or outer sheath cells are either

translucent, without chloroplasts, or they may

contain green pigment in plastids similar to, or

differing from, the cholorplasts of the chlorenchyma

of the mesophyll. Many early workers ignored the

contents of these sheath cells and represented them as

being empty (Lewton-Brain, 1904; Lohauss, 1905;

Prat, 1936; Vickery, 1936). However, the nature of

these contents, whether typical chloroplasts or

specialized plastids is undoubtedly a character of

systematic importance as Brown (1958) correctly

stresses. In this connection, it is important that in

older leaves of wheat few or no plastids are present

in the bundle sheaths (Percival, 1929). Thus, age of

leaf sampled must be standardized for this character

to be reliable. The detailed structure and function

of these bundle sheath cells will be elaborated on

later.

The arrangement of the chloroplasts in the bundle

sheath cells has been used for diagnostic purposes —

whether they display a horse-shoe shaped arrangement

against the inner or outer tangential wall, or an even

distribution in the cells. However, Roth (1968), has

shown in xerophytic species such as Sporobolus

virginicus that the horse-shoe shaped arrangements

occur under low-intensity illumination, such as in

the mornings, afternoons, at night and in rolled up

leaves, while on sunny days the chloroplasts are

evenly arranged all over the walls. This is probably an

adaptation to efficient light utilization, but cannot

carry much weight as a diagnostic character.

The single or outer sheath may possess girder-like

extensions which are of diagnostic importance. The

extensions either consist of colourless cells resembling

those of the bundle sheath or there may be a gradual

transition to the sclerenchyma fibres of the adjacent

strand. The extensions may be uni-seriate to very wide,

and sometimes flank narrow sclerenchyma strands or

girders as in the first order bundles of Garnotia

scoparia (Tateoka, 1958). Burbidge (1946) and Decker

(1964) discuss the unique outer bundle sheath of

Triodia leaves where the sheath of one bundle is

continuous with those of adjacent bundles.

The adjacent sclerenchyma girders may interrupt

the single, or outer, sheath to a greater or lesser

extent, either adaxially, abaxially or both. Thus, the

outer or single bundle sheath may be entire, horse-shoe

shaped or even reduced to two lateral arcs. In

Arundinella leptochloa (Tateoka, 1958) this sheath is

only developed adaxially adjacent to the xylem, not

being replaced by a sclerenchyma girder abutting on

the lateral parts of the phloem. |

Schweickerdt (1941) has used the number of cells i

comprising this sheath to distinguish the closely

related genera of Monelytrum and Tragus. This cell

number appears to exhibit excessive variability in

most instances, but it is relatively constant for the

third order bundles of groups such as the Paniceae. !

Various degrees of possible reduction of the vascular

bundles, as perhaps evidenced by relic bundle sheath

cells have been observed in Arundinella (Vickery, 1935)

and in Garnotia (Tateoka, 1958). In these grasses the

first order vascular bundles have well developed outer

sheaths, Garnotia even having abaxial extensions. ii

In smaller bundles these extensions are reduced, and

only a few cells, resembling those of the bundle lj#

R. P. ELLIS

101

sheath, are irregularly present on the sclerenchyma

girders or strands. The extreme situation is found

between successive vascular bundles of some Arun-

dinella and Garnotia species, where solitary, or groups

of two to six cells, similar to those of the bundle

sheaths are present, scattered in the mesophyll.

Tateoka (1958) calls these groups of sheath-like cells,

lacking associated vascular tissue, distinctive cells.

Carolin, Jacobs & Vesk (1973) have shown that these

distinctive cells are in fact isolated files of parenchyma

sheath cells connecting with the bundle sheath proper.

They consider this to be a specialization to the wide

expanses of mesophyll between adjacent vascular

bundles found in these species. These distinctive cells

ensure that all mesophyll cells are ultimately in

contact with the bundle sheath parenchyma cells.

The inner sheath normally consists of living, thick-

walled, chloroplast-free cells which are elongated

parallel to the vascular bundles. In section the cells

of this sheath are nearly always smaller in diameter

than the outer sheath and possess thickened walls,

the thickening often being more conspicuous on the

radial and inner tangential walls. Exceptions to these

generalizations do occur as in Aristida, Elytrophorus

(Schweickerdt, 1942) which has a thin-walled inner

bundle sheath, and the Meliceae (Decker, 1964)

where the inner sheath cells are uniformly thickened.

This sheath often resembles an endodermis and it

has been shown to have similar physiological activity

(van Fleet, 1942). It has been variously called the

endodermis or mestome sheath in the past. Unfor-

tunately, the term mestome sheath, has been used for

parenchymatous sheaths as well and modern authors

(Metcalfe, 1960; Barnard, 1964), therefore, refer to it

merely as the inner sheath.

The inner sheath is sometimes difficult to recognise,

especially in cases where the cells are not distinct

from those of the fibres of the vascular bundle ground

tissue. However, cells of this layer are always in

contact with the metaxylem vessels (Vickery, 1935).

Thus, if large inflated cells are immediately adjacent

to the metaxylem vessels it can be assumed that no

inner sheath is present.

In addition to sometimes not being sharply

differentiated from the bundle ground tissue, in some

smaller bundles the inner bundle sheath is sometimes

only developed adjacent to the phloem (Fahn, 1967).

For this reason Metcalfe (I960) proposed the intro-

duction of a third bundle sheath category — inter-

l mediate — in addition to single and double sheaths.

1 The inner bundle sheath cells of Aristida are

anomalous in two ways: the diameter of the inner

bundle sheath cells is greater than those of the outer

sheath, and they contain numerous chloroplasts.

These larger inner sheath cells may also have

uniformly thick walls. Holm (1901) was the first to

record this double bundle sheath, Vickery (1935)

interpreted this condition correctly and Lommasson

(1957) considered it to be of taxonomic importance,

so much so as to warrant the placing of Aristida in a

separate tribe. Theron (1936) attached little

taxonomic importance to the sheaths of Aristida ,

probably having misinterpreted the specialized

bundle sheath.

Lommasson (1957) was further able to show that, in

the leaf sheath, where both bundle sheaths lack

chlorophyll, on the adaxial side against the culm,

the cells of the outer bundle sheath were larger than

those of the inner bundle sheath. The transition to

the opposite condition, as in the leaf blade, occurs

where chlorophyll is formed in the bundle sheath

cells and mesophyll. He advanced the explanation

that this anomalous condition represents the result

of special activities occurring at the chlorophyll —

vascular interface.

The structure and function of the parenchyma cells

of the outer, or single sheath, and their dimorphic

chloroplasts have been extensively studied. Zirkle

(1929) recognised that the cells of the bundle sheath

in Zea contained specialized chloroplasts concerned

with starch storage. In 1944, Rhoades and Carvalho

reported that in panicoid grasses, such as Zea and

Sorghum, the single parenchyma sheath contained

large specialized plastids, functioning in starch

formation and storage. In the outer parenchyma

sheath cells of the festucoid grasses, as represented by

barley, wheat and oats, they noticed the plastids to be

smaller than those in the mesophyll. In these species

the plastids of the chlorenchyma, as well as those of

the bundle sheath cells, form starch.

Rhoades and Carvalho further demonstrated that

in festucoid grasses the mesophyll cells photosyn-

thesise and accumulate starch during daylight. In

these grasses the outer parenchyma sheath cells have

numerous plastids which, however, are smaller than

those of the mesophyll. Small amounts of starch,

lying in the central region of the plastid were found

in the sheath plastids as well as those of the mesophyll.

In Zea and Sorghum the chlorenchyma cells

photosynthesise, but do not store starch at all.

Thus, no trace of starch was found in the chloroplasts

even if abundant starch was present in the bundle

sheath plastids. The plastids of the single sheath

appear to elaborate, and temporarily store starch,

but have little photosynthetic activity. These workers

showed, that the starch found in these plastids was

derived from soluble carbohydrates made in the

mesophyll plastids, and translocated to the sheath

cells where starch synthesis occurred, and not from

sugars synthesised in the plastids of the bundle sheath,

which could be capable of photosynthesis since they

contain a green pigment.

This transformation of soluble carbohydrates to

starch in the bundle sheath plastids occurs only when

the rate of movement of sugars into these cells is

greater than the translocation from the bundle sheath

cells into the vascular bundles. Starch, which

accumulates throughout the day, is transformed back

into soluble carbohydrates during the night, so that

by morning the bundle sheath plastids are devoid of

starch. Roth (1968) considers the possibility that these

cells function in water storage as well, especially in

xerophytic species.

Grasses with this specialized type of bundle sheath

structure and function have been termed the eupani-

coid subtype of the panicoid subfamily (Brown, 1961).

The other subtype, the chloridoid grasses have a dark

green parenchymatous sheath and a single layer of

pale green radial chlorenchyma. Here the sheath cells

photosynthesise in addition to accumulating starch.

Brown (1961) postulates that the chlorenchyma may

have assumed some unknown function to replace the

activities of photosynthesis and starch storage which

it has more or less lost. Otieno (1967) is of the opinion

that the radiating mesophyll cell layer, of these

chloridoid grasses, constitutes part of the bundle

sheath which is then composed of three layers.

This is not acceptable because all intermediate types

between irregular and radiate chlorenchyma are

found.

The starch-free bundle sheath plastid has numerous,

small colourless areas, resembling vacuoles, contained

within a peripheral rim. It is in these vacuolar-like

regions that starch is deposited. Thus, each plastid

102

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

contains numerous simple starch grains embedded

in, but protruding from, the surface of the plastid.

Ultrastructural studies of these dimorphic chloro'

plasts of the panicoid grasses, by Laetsch & Price

(1969) on Saccharum, Downton & Pyliotis (1971) on

Sorghum bicolor and Andersen et al. (1972) on Zea

show, that the bundle sheath chloroplasts loose their

grana during ontogeny. Young bundle sheath chloro-

plasts have well-developed grana, but mature plastids

are agranal whereas mesophyll chloroplasts contain

grana irrespective of their developmental stage. They

conclude that the structure of the specialized chloro-

plasts in bundle sheath cells is a result of reduction

and that this chloroplast dimorphism is a

specialization of labour. There are various degrees of

reduction in the sheath chloroplasts. Zea, Coix and

various members of the Paniceae and Andropogo-

neae have rudimentary grana and Carolin, Jacobs &

Vesk (1973) have shown that the Eragrostoideae in

fact have well-developed grana. Johnson (1964)

suggests the possibility that the degree of specialization

of these chloroplasts has phylogenetic significance.

The chloroplast-containing sheath cells contain,

in addition, a remarkably high concentration of

additional cellular organelles, such as mitochondria,

endoplasmic reticulum and perioxisomes, but vacuoles

are difficult to locate. The differences in the ultra-

structural features of the cells of the bundle sheath and

those of the chlorenchyma further suggest the existence

of differences in the functions of these cells (Doby-

china, 1970; Carolin, Jacobs & Vesk, 1973).

This detailed discussion of the chloroplasts and the

bundle sheath has been considered necessary because

in recent years this aspect has received considerable

attention from the point of view of the Kranz syn-

drome. Thus, grasses with bundle sheaths containing

specialized chloroplasts are typical of the high photo-

synthethic capacity grasses and those with no, or

normal chloroplasts are characteristic of low photo-

synthetic capacity grasses (Black, 1971).

At present the advantages of highly developed

sheath cells still remain unclear. Black (1971) infers

that the anatomy of the high photosynthetic capacity

plants results in more rapid rates of translocation,

and perhaps higher concentrations of translocates,

which may both prevent a feedback type of inhibition

of photosynthesis by a product such as starch.

Nutrients may, in addition, be supplied to non-

photosynthetic parts of the plant such that high growth

rates are facilitated.

The possibility of the inner bundle sheath, or the

single sheath, when the inner sheath is lacking,

having functions and characteristics similar to an

endodermis has received considerable attention in the

literature. In grasses with a single bundle sheath,

the cells of this sheath do not have the anatomical

characteristics associated with an endodermis, such

as Casparian strips, thicker radial and inner tangential

walls or suberised walls. This is applicable to a

number of inner sheaths as well.

Van Fleet (1950) demonstrated, with a variety of

histochemical reactions, however, that the cells of

these sheaths share common substance reactions and

may be induced to develop typical endodermal

characteristics. Thus, normally the single sheath cells

contain chlorplasts and do not exhibit characteristics

of an endodermis. But in variegated and albino leaves,

or in etiolated leaves. Van Fleet showed that those

areas lacking chlorophyll show characters of a typical

inner bundle sheath, i.e. endodermal characteristics.

In Pennisetum villosum and Oplismenus hirtellus he

further showed that casparian deposits are developed

on the radial walls prior to the unilateral desposition

characteristic of the mature inner sheath. Endodermal

characteristics, therefore, appear at varying stages

in the development of the leaf in at least some of the

cells of an atypical inner sheath. Thus, in any sheath

bounding the vascular tissue, some or all, attributes of

a true endodermis can be detected. Van Fleet points

out that the endodermal characteristics are always

more pronounced opposite the phloem than opposite

the xylem. Once again for comparative work it is

essential to standardize upon leaves of equivalent

developmental stages.

Schwendener (1890) stated that the walls of the

parenchymatous sheath of Zea contained suberized

lamellae, something Van Fleet (1950) was not able to

show for normal maize leaves. O’Brien & Carr (1970)

also described a suberized lamella, which is probably

the site of suberin deposition, in the walls of the

inner sheath of Triticum and Avena as well as in the

parenchyma sheath of maize. The suberized lamellae

modify the plasmodesmata of the numerous pit-fields

connecting the cells of the inner sheath with those of

the outer sheath and vascular parenchyma, or between

cells of the single parenchyma sheath.

Using these ultrastructural details O’Brien & Carr

(1970) propose a function for the inner bundle

sheath and a further function, in addition to photo-

synthesis and starch storage, for the single

sheath. These workers are of the opinoin that the

suberized lamellae are relatively impermeable to

water and thus there is the possibility that in grass

leaves water loss is regulated at the vascular bundle

as well as by the stomata. This control of water loss

at the vascular bundles is probably essential for an

adequate supply of solute to be maintained to the

leaf tip, because, under conditions of water stress

the lower part of the blade could conceivably transpire

all the water available to the leaf. If the suberized

lamellae restrict passive loss of water from the

vascular bundles to the mesophyll, forcing the water

to follow a symplastic route through the sheath cells,

the water flow across the sheath could be regulated

by the activity of the sheath cells.

Further circumstantial evidence for this possibility

is that transfer cells, which are believed to help regulate

solute exchange between tissue systems, are absent

from the leaves of grasses. In dicotyledon leaves these

transfer cells are thought to assist exchange between

xylem and phloem, and between these and the

adjacent mesophyll. O’Brien & Carr reason that if

passive loss of water is restricted from the vascular

bundles because of the suberized lamellae, and if the

bulk of solute transfer must pass through the plas-

modesmata, then transfer cells are unnecessary.

Sclerenchyma of the leaf

In the grass leaf blade, the sclerenchymatous tissue, |

or the sterome, includes all fibres as well as other

thick-walled cells in certain instances. The scleren- i

chyma is commonly found in association with the

vascular bundles, with the midrib or keel and in the

margin.

Xylem and phloem fibres, and thickened vascular

parenchyma, constitute part of the ground tissue of

the vascular bundles. The mestome, or inner bundle

sheath, is also fibrous, the fibres appearing, in some

instances, ontogenetically the same as those of the

hypodermal sclerenchyma, but differing in that they

possess numerous pits which are lacking in the true

R. P. ELLIS

103

sterome (Artswager, 1925). This tissue is included

in the description of vascular bundle structure and

will not be discussed further here.

The sclerenchyma associated with the vascular

bundles is in the form of sub-epidermal longitudinal

bands following the course of each vascular bundle.

In transverse section this circumvascular sclerenchyma

may surround, be in contact with, or may be situated

above or below, but not connected to the bundle or

it’s sheath. This sclerenchyma tissue is termed a

strand when it does not extend sufficiently deeply into

the mesophyll to make contact with the bundle

sheath cells (Metcalfe, 1960). These strands, therefore,

appear in section as small hypodermal “islands” of

thickened tissue situated above and below, or on

one side only of each vascular bundle. When the

inner face of a group of fibres is in contact with,

disrupts, or envelopes the bundle sheath, it is termed

a girder (Metcalfe, 1960). It resembles a girder in

transverse section, extending from either or each

epidermis to the bundle sheath. When girders are

continuous from the vascular bundle to the epidermis

on either side it is termed an I-beam construction

(Gould, 1968).

All distribution patterns of the circumvascular

sclerenchyma occur either as strands or girders. In

addition, preliminary observations in the present

study suggest that a further distinction between

girders in contact with, and those disrupting the

parenchymatous sheath, may be useful taxonomically.

In leaves possessing both strands and girders, the

former are usually associated with the third order

vascular bundles and the latter with larger bundles.

Other leaves may have strands or girders only

developed in association with all the vascular bundles

of the leaf. The distribution and arrangement of the

sclerenchyma associated with the bundles is, therefore,

useful diagnostically. Furthermore, the amount of

sclerenchyma present varies from species to species

(Metcalfe, 1960). This can be misleading, however,

because marked intraspecific variation will be found

in specimens from different localitities or from

different seasons as the work of Burduja & Toma

(1970) on Deschampsia flexuosa and other species

has shown. They found that on flowering tillers there

was less sclerenchyma in the leaves than on non-

1 flowering tillers where the fibre walls were also thicker

with the lumen smaller. Taxonomically sclerenchyma

distribution is seldom of more than specific diagnostic

| importance.

The distribution of sclerenchyma associated with

the vascular bundles can be correlated with ecological

factors as well. Grasses from arid areas are thus

characterized by well developed sclerenchyma tissue

while many tropical grasses often have a high

proportion of the smaller bundles not accompanied

by sclerenchyma.

The function of all the sclerenchyma present in the

leaf is undoubtedly to provide mechanical support

for the softer tissues. The development of scleren-

chymatous tissues makes possible the withstanding of

the physical stresses and strains imposed on an

elongated, straplike leaf such as is predominant in the

Poaceae. An additional function of the fibres is seen

by their frequent silicification, especially in older

leaves (Parry & Smithson, 1964) and they, therefore,

can act as a depository for excessive silica. Silicified

fibres result in elongated, pointed needles of silica.

The thickening of the fibre cell walls is usually by

lignification, but in many specimens the staining

reaction varies across a single sclerenchyma girder,

or in girders from different parts of the lamina. In

these cases, when stained with safranin and fast green,

there is a gradation from typically red-staining fibres to

structurally similar fibres which attract the blue or

green dye. Hoefer (1941-42) also found variations

in the reaction of fibres in different parts of the leaf

of Stipa tenacissimci when treated with lignin stains.

In addition, lignin tests with phloroglucin and hydro-

chloric acid and tests with Maules’s reagent did not

give identical results. Hoefer’s tests showed, in addition

that there is zonation in the degree of lignification in

different layers of the cell wall of individual fibres as

seen with transverse section. It appears, therefore, that

the degree of lignification varies, but in addition,

lignins of various chemical constituents appear at

present to be grouped under the general term

“lignin”.

In certain grasses sub-epidermal longitudinal

strands are found between successive vascular bundles,

either alone or in addition to the circumvascular

strands or girders. These supernumary strands are

usually abaxial and situated opposite the bulliform

cell groups or furrows (Lewton-Brain, 1904). When

sclerenchyma is found between the bundles in

expanded leaves it is of special importance diagnos-

tically.

Continuous abaxial hypodermal bands are found

in certain species, especially those with acicular

leaves. These bands may be a regular, narrow strip,

2-4 cells deep, located immediately beneath the

epidermis, or may result from lateral extensions of

the fibrous tissue comprising strands or girders.

Mechanical tissue of the leaf margin may occur in

the form of a cap or hood. When the marginal

sclerenchyma is not in contact with the lateral bundle,

it is termed a cap, and a hood is formed when this

sclerenchyma extends above or below the lateral

bundles. The fibres of the hood may, or may not, be

in contact with the bundle sheath of the ultimate

bundle, but always extend inwards from the margin,

as far as, or further than, this bundle. This lateral

fusion with the sterome tissue associated with the

lateral bundles forms the hood-like structure

(Goossens, 1938). In some instances, there is no

sclerenchyma development at the extreme margin but

the ultimate and penultimate lateral bundles may be

intimately associated with specialized sclerenchyma

development. Immediately interior to the fibrous

cap may be situated normal mesophyll cells, small

groups of enlarged, colourless parenchyma cells or a

lateral intercellular duct may be developed.

Sclerenchymatous tissue is also found developed

in association with midribs and keels. This has been

described together with the median vascular bundles,

midribs and keels.

In reduced, permanently infolded leaves the

distribution of the sclerenchyma follows that outlined

above in basic pattern, but appears symmetrical

and distinctive because of the nature of the leaf

blade. In these leaves this arrangement of the

mechanical tissue is of taxonomic importance because,

in assuming the permanently infolded form, the

leaves of different grasses have not acquired precisely

the same sclerenchyma tissue arrangements (Metcalfe,

1960).

Mesophyll

In the Poaceae, the term mesophyll is generally

applied to the ground tissue occupying all the space

in the leaf not occupied by the vascular bundles, the

bundle sheaths and the sclerenchyma. The mesophyll

can be subdivided into the assimilatory chlorenchyma

104

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

and the colourless parenchyma which consists of

translucent cells often in close association with the

bulliform cells. For this reason further details of

colourless parenchyma will also be found under the

discussion of bulliform cells.

Grass leaf blade chlorenchyma seldom exhibits a

distinct differentiation into pallisade and spongy

regions as seen in transverse section. In some species

the adaxial chlorenchyma cells are more regularly

and vertically arranged than the remainder. Metcalfe

(1960) correctly stresses that this distinction is rare

and at best unclear. Statements in the literature

often imply that there is greater contrast between

these zones than there actually is.

In dicotyledons, as Watson (1942) has shown with

Hedera helix, the English ivy, the leaves respond to

strong light by the production of pallisade tissue.

On the same plant shade-grown leaves have no

pallisade tissue at all. These cell differences are caused

by factors affecting the processes of vacuolation and

enlargement of the cells. Thus, the uppermost layer

of mesophyll cells has an increased osmotic value due

to a change in the starch-sugar ratio. This, according

to Watson (1942), results in the absorbtion of water,

increased vacuolation and expansion, with the

resultant formation of pallisade tissue.

In the Poaceae, the above does not appear to apply

and in certain instances the opposite may be the

case. In the genus Panicum, the shade-loving forest

species often have the upper layer of chloren-

chymatous cells more or less vertically arranged in

a pallisade manner in contrast to most other species

of this genus where the assimilatory tissue is more

of less radiate.

The arrangement of the chlorenchyma cells appears

to be of fundamental taxonomic significance. Thus,

in the festucoid type, the chlorenchymatous tissue

is more or less homogenous without being arranged

in any definite pattern in relation to the vascular

bundles. In grasses of the panicoid type the assimi-

latory cells are arranged in a radiating manner around

the vascular bundles as seen in transverse section.

Each bundle is, therefore, situated in the centre of a

regular circle, or partial circle of chlorenchyma.

The former irregular arrangement is, in addition,

generally associated with leaves in which the vascular

bundles are widely spaced, whereas the radiate

condition and closely placed vascular bundles are

correlated (Vickery, 1935).

The division of the Poaceae into two sub-families

with radiate or non radiate chlorenchyma is an

oversimplification. Brown (1958) shows that within

the panicoid grasses the Chlorideae and related

tribes have the regularly radiate condition, whereas

this radiate condition is less regular in other panicoid

genera such as Andropogon and Panicum. Thus, the

radiate condition is of more than one kind, and

grasses with partially or incompletely radiate chloren-

chyma also occur, as Metcalfe (1960) stresses.

Nevertheless, the distinction between radiate and

non-radiate chlorenchyma is important taxonomically

provided that the limitations are recognized. For

example, both conditions may occur in the same

genus, as in Sporobolus where many species have

regular, elongated chlorenchyma cells radiately

arranged around the bundles, but in S. panicoides

the cells are irregular in shape and arrangement

(Goossens, 1938).

Early workers correlated the non-radiate condition

with grasses from temperate regions, whereas radiate

assimilatory tissue was linked with tropical grasses.

The geographical pattern is not so distinct, however,

with many exceptions and Metcalfe (1960) remarks

that the mesophyll arrangement is of more funda-

mental significance taxonomically than geographically, j

In transverse section the individual chlorenchyma

cells of most panicoid grasses appear elongated,

narrow and tabular in shape. Their long axes are at

right angles to the bundles. In festucoid grasses the

cells are often irregular in shape and size. In some

species, such as Poa annua, the largest cells are found

nearest the vascular bundles (Bobrov, 1955) or the

chlorenchyma cells abutting on both epidermides

may be somewhat elongated and regular as in

Triticum vulgar e (Hayward, 1948). The irregular

condition may be characterized by small, isodia-

metric cells tightly packed together or by irregularly

shaped cells with many intercellular air spaces between

them. The largest intercellular air spaces are commonly

seen subtending the stomata and projecting deeply

into the mesophyll. Pool (1923) illustrates such

distinct sub-stomatal chambers in Andropogon

furcatus.

Slade (1970) in a study of sun and shade-grown

leaves of Poa alpina has shown that in sun leaves the

mesophyll cells are more or less isodiametric and

not longitudinally elongated as in shade leaves. The

intercellular spaces are relatively inconspicuous in

sun leaves but extensive in shade leaves as seen in

longitudinal section. Slade also demonstrated that in

transverse section there was no noticeable change in

the horizontal widths of the intercostal zones of

chlorenchyma between adjacent vascular bundles, but

that the depth or thickness of the mesophyll was

greatest in sun-grown leaves. This is as a result of

reduction in cell size and intercellular spaces in shade

leaves seen in transverse section in contrast to the

situation in longitudinal section above.

The overall effect of these anatomical changes in

sun and shade leaves is to physically weaken the leaf

blade. Slade (1970) has shown, in addition, that in low

light intensity the cell walls are thinner and a well

defined cuticle may be absent.

It appears, therefore, that the light intensity under

which a leaf develops has a profound effect on the

arrangement of the assimilatory cells into pallisade

and spongy mesophyll, especially in dicotyledons

(Watson, 1942), as well as the size and shape of the

individual cells as shown by Slade (1970). The

applicability of these findings to the chlorenchyma

cell structure and arrangement in the Poaceae as a

whole does not appear clear at present. Thus, if the

radiate and non-radiate conditions are of funda-

mental phylogenetic significance it appears unlikely

that major differences of tissue arrangement will occur

under varying light intensities. Some generalizations

do seem to apply and most forest dwelling species

examined are characterized by very thin mesophyl

tissue. Prosphytochloa prehensilis is an extreme

example with the whole leaf being only four cell

layers thick (de Winter, 1951).

Intercellular air spaces occur between the chloren-

chyma cells of many grasses belonging to tribes of

both the major subfamilies. These spaces may be very

conspicuous in hygrophilous species (Arber, 1934;

Vickery, 1935). In other aquatic species there are, in

addition, distinct air cavities or lacunae present.

All the species of Elytrophorus have these lacunae

situated between adjacent vascular bundles

(Schweickerdt, 1942) and in Vetiveria they occur over

the smaller bundles (Kanmathy, 1969). The lacunae

are traversed by colourless aerenchyma cells which are

R. P. ELLIS

105

often stellate in shape and represent diaphragms in the

air cavities. Kanmathy (1969) reports sclerotic

strands interspersed in the stellate cells.

Cavities developing as a result of breakdown of

parenchymatous tissue, especially in the region of

the keel, and in the keel itself, are found in many

grasses. These differ from lacunae in being indistinct

with the breakdown of the cells continuing. This type

of cavity has been used diagnostically for Molinia

caerula (Lewton-Brain, 1904) and Sporobolus artus

var. lysigenatous (Goossens, 1938).

A further modification of the mesophyll tissue is

found in certain markedly infolded leaves where the

chlorenchyma is confined to small bands adjacent to

the adaxial and abaxial cleft-like furrows which

contain the stomata. The tissue between the groups of

assimilatory tissue is made up of colourless

parenchyma. This unique type of anatomy is dis-

cussed by Burbidge (1946, 1946a) for Triodia species.

Burbidge believes that this reduction of the chloren-

chymatous tissue results in a corresponding narrowing

of the width of the furrows and consequent pro-

tection of the stomata. De Wet (1956) found localized

bands of chlorenchyma cells around the abaxial

grooves in species of Merxmuellera ( =Danthonia )

and suggests that this may indicate relationships.

In some species, groups or isolated, bundle sheath-

like cells with chloroplasts, but lacking associated

vascular tissue are found scattered in the mesophyll.

These are termed distinctive cells (Tateoka, 1958)

and are described under vascular bundle sheaths.

Bamboos, and a few other grasses, are characterized

by the presence of fusoid cells in the mesophyll of

their leaves. These cells have been termed enlarged

parenchyma cells (Page, 1947) and in transverse

section have a fusiform or pyriform outline, being

elongated transversely and alternating with chloren-

chyma cells and the vascular bundles. Adjacent fusoid

cells may be separated by a single vertical column of

chlorenchyma or many chlorenchyma cells may

separate them. The bulliform cells are located imme-

diately adaxially to these chlorenchyma cells

separating adjacent fusoid cells.

These conspicuous colourless cells have been

incorrectly interpreted as intercellular spaces

(Jacques-Felix, 1955). Their cellular nature was

confirmed by Page (1947) and Metcalfe (1956).

The fusoid cells are, in fact, narrow and plate-like

with the long axis of the cells lying at right angles to

the long axis of the lamina. Longitudinal sections of

the blade show these cells to have very narrow

lumina in transverse section, the cells having collapsed

in such a way that the tissue resembles a row of

“I’s”. Between the dead, collapsed, mature cells

there are large spaces which connect with the inter-

cellular spaces of the chlorenchyma. In certain species

the fusoid cells fail to collapse, but become rounded

and separated from one another. The regularity and

form of these collapsed cells seems to indicate that

some force operates on the whole leaf at the same time

(Page, 1947).

In addition, the Bambusae, as well as the Oryzeae,

are characterized by having chlorenchyma com-

prised of what, in the grass anatomy literature, have

become known as arm cells. These have also been

called irregularly lobed or cleft cells (Page, 1947) or

plicate mesophyll (Esau, 1960). These so-called arm

cells possess inwardly directed projections or folds

which may be continuous across the cell as seen in

section and completely divide the cells into elongated

compartments. The infoldings may end blindly in the

lumina as well. These projections may be developed

from the upper or the lower wall of the chlorenchyma

cell, or from both the upper and lower, or from all

walls (Brandis, 1907).

Chih-Ying Wu (1958) has shown that in some

bamboo species the arm cells located above the

fusoid cells have projections from the lower walls,

whereas the cells below the fusoid cells and cavities

have folds projecting from the upper walls. This

phenomenon is mentioned by Arber (1934), but in

the Oryzeae, even when fusoid cells are present, the

arm cells illustrated by Tateoka (1963) have infoldings

from all directions and of equal length.

It is not certain how these projections arise. They

could be infoldings of the cell walls as an illustration of

Carolin, Jacobs & Vesk (1973) indicates, but in some

grasses their appearance suggests that they arise as fine

partitions that at first traverse the cell completely,

but subsequently become broken as the cells become

enlarged (Metcalfe, 1960). Haberlandt (1884) called

these “armpalisadezellen” and explained the signi-

ficance of the infoldings as increasing the inner

surface area of the cell, thereby creating space for

more chloroplasts. Infoldings are common in many

grass genera, and are termed arm cells or peg cells, and

in the bamboos and their relatives it is possible that

the pegs become “fused”, eliminating the air spaces.

Fusoid and arm cells are important taxonomic

characters. Thus, fusoid cells are especially charac-

teristic of the Bambusae (Metcalfe, 1960), and when

they occur in other genera, such as Oryza (Tateoka,

1963), affinities between these genera and the

bamboos may be indicated. Arm cells, although

characteristic of the Bambusae and Oryzeae, may

also show variations in their occurrence. Thus, Oryza

tisseranti is exceptional amongst the Oryzeae in not

having arm cells (Jacques-Felix, 1958). Gordon-

Gray & Ward (1971) noticed mesophyll cells with

invaginated walls in Phragmites and suggest that

this may reflect closer relationships to the Bambusae

and Oryzeae than was previously thought possible.

The extreme case is that of Saccharum, where Merida

(1970) found cells with internal folds situated below

the stomata in some varieties. This apparent

occurrence of arm cells in a member of the Andro-

pogoneae is certainly surprising.

Bambusoid and oryzoid grasses are very poorly

represented in South Africa. For this reason, the

accompanying descriptive key is not concerned w ith

the details of arm and fusoid cells to any great degree.

As presently constituted it cannot be expected to

differentiate between the various bamboo genera.

The colourless parenchyma constituent of the

mesophyll, especially that intimately associated with

the bulliform cells is described, together with these

bulliform cells, under the description of bulliform

cells.

Bulliform and colourless cells

Bulliform cells are single, translucent cells, or

groups of colourless cells, constituting part of, or

the entire epidermis, but differing from other epidermal

cells in being larger and more inflated. These cells

occur most commonly, but not exclusively, at the

bases of adaxial furrows (Metcalfe, I960). Shields

(1951) restricts the term bulliform cell to inflated

epidermal cells present in the adaxial furrows.

However, structurally similar cells may be present in

the abaxial epidermis as well as, or instead of, the

adaxial epidermis. Other terms used to describe

these cells are hinge cells and motor cells, but the

106

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

descriptive term bulliform, denoting their inflated

appearance, is preferred to these terms referring

to their disputed function in leaf involution.

Bulliform cells often form longitudinal, parallel,

intercostal bands, and are commonly associated with

underlying groups of structurally similar cells, which

usually form vertically elongated units in the meso-

phyll when viewed in transverse section. Shields (1951)

designated these associated cells, hinge cells. Here

they are grouped with the colourless cells, a term

incorporating all the translucent cells present in

the mesophyll excluding bundle sheath cells devoid

of chloroplasts, fusoid cells of bamboos and

epidermal cells. The bulliform and associated

colourless cells may appear in transverse section as

deep girders, or may be restricted and superficial.

The colourless cells associated with the bulliform

cells are variable in shape and size, but all colourless

cells, whether associated with the bulliform cells or

not, are always without chloroplasts.

Bulliform cells have long been recognised as

valuable taxonomic characters. As early as 1907,

Brandis reviewed the distribution of bulliform cells

in the Poaceae as a whole. The major character

subdivisions he used were the presence of bulliform

cells in the adaxial epidermis only, the presence of

bulliform cells in both adaxial and abaxial epidermides,

and the absence of distinct bulliform groups. The

adaxial bulliform cell distribution was further sub-

divided into those grasses where bulliform cell groups

alternated with all vascular bundles, those where the

bulliform cells were present between successive first

order bundles, but located over the third order

bundles, those with bulliform cells on either side of

the midrib and at the margins, those with bulliform

cells only located on either side, or above the midrib.

Using these distributional criteria of bulliform cells

Brandis (1907) found uniform distribution in the tribes

Bambuseae and Maydeae only. In all the other

tribes studied various genera had different bulliform

cell arrangement in some of their species.

The arrangement, frequency, distribution, as well as

the relative size and shape of the bulliform cells and

colourless cells are of taxonomic importance especially

at the specific level. Thus, in Elytrophorus,

Schweickerdt (1942) found the presence of conspicuous

bulliform cells flanking the midribs only, to be the

the oustanding diagnostic character of E. africcmus,

whereas in E. interruptus bulliform groups are

present throughout the leaf width, but decrease in

size towards the margin. Furthermore, in one species,

the shape of the individual bulliform cells in trans-

verse sections is rectangular, whereas in the other

species they are trapezoidal with the outer wall being

shorter than the inner wall. In Leersia hexandra

there are groups of bulliform cells between all the

vascular bundles and on both epidermides but other

Leersia species have only two groups of bulliform

cells on either side of the midrib (Holm, 1895).

Goossens & Theron (1934) found the bulliform cells

to be of significance in classifying the different

varieties of Themeda iriandra. The arrangement of

the colourless cells in relation to the bulliform cells

can be of specific diagnostic importance. In Imperata

cylindrica (Vickery, 1935), where the bulliform cell

groups occur over third order bundles, two rows of

colourless cells are produced towards the abaxial

surface, one on either side of each bundle.

The function of the bulliform cells has been the

subject of much controversy in the literature and a

totally satisfactory explanation of their function has,

as yet, not been forthcoming. Duval-Jouve (1875) was

the first to describe these cells and believed that the

opening and inrolling of the blade resulted from

changes in the turgor pressures of these cells. Some

workers, such as Breakwell (1915), have carried this

even further by postulating that rolling occurs only

at times of water shortage and results in the leaf

becoming tubular, thus protecting the greater number

of adaxial stomata from excessive transpiration. This

teleological viewpoint is not supported by the fact

that many grasses, including drought resistant types

which exhibit inrolling, have more stomata on the

exposed abaxial surface than the protected adaxial

surface (Vickery, 1935). Hayward (1948) mistakenly

states that the outer surfaces of the bulliform cells

are not cutinised, and thus readily lose water through

diffusion. Metcalfe (1960) points out that in many

grasses exhibiting involution the outer wall of the

bulliform cells may be considerably thickened.

The conventional explanation of bulliform function

thus accepts that the bulliform cells regulate the

movement of the blade when it opens and closes.

Goossens & Theron (1934) draw a comparison

between bulliform cells and stomatal guard cells and

the changing of cell shape and turgidity with turgor

changes. However, whether the bulliform cells

actually cause the rolling movements by collapse due

to water loss, or whether their size and plasticity

merely permit them to be compressed and so allow

rolling to occur, has been the subject of some con-

troversy.

Numerous facts point to caution in explaning the

function of these cells. Brandis (1907) remarks on the

fact that in mature leaves the bulliform cells may

become entirely filled with solid silica making the

leaf rigid. Bor (1960) quotes Goebel (1926) as

describing nyctinastic movements in Leersia hexandra

in which the leaves fold up in dull weather and

flatten again in full sunlight. These movements

obviously are not dependant on different water

regimes within the bulliform cells. In addition,

involution may occur in the absence of bulliform cells

and often the thin lateral walls of bulliform cells are I

not collapsed or distorted in naturally rolled or

wilted grass leaves (Shields, 1951). j

Tschirch (1882) challenged this theory that move-

ments depended only on turgor changes in the bulli-

form cells. While he agreed that rolling and unrolling

of grass leaves was influenced by the loss and uptake

of water, he thought that these movements were due,

at least in some cases, to changes in the leaf fibres

rather than the epidermal bulliform cells. He explained

that the adaxial, subepidermal schlerenchyma fibres

have a high capacity for imbibition and a marked

tendency to shrink when dehydrated. This results in

contraction of the adaxial ribs causing inrolling

because the inflexible lower epidermis, reinforced by

sclerenchyma gives the abaxial surface an unbroken

rigidity. This contraction, coupled with cohesion

among the chlorenchyma cells and among the bulliform

cells may result in leaf involution. Thus, involution

has also been defined as a cohesion and shrinkage

phenomenon, the result of a differential contraction

of the mesophyll and the rigid lower epidermis. As

the mesophyll cells shorten in drying, they become

stretched around the vascular bundles, causing the

leaf to roll.

Shields (1951), in a detailed study of grass leaf

involution, has shown that differential shrinkage of

the adaxial and abaxial rib surfaces occurs. This

shrinkage was more pronounced in all the species

studied on the adaxial rib surface. This illustrates the

lateral shrinkage in the adaxial sclerenchymatous

R. P. ELLIS

107

fibres, where present, and in the rnesophyll. In leaves

lacking adaxial sclerenchyma girders or strands,

large adaxial rnesophyll cells collapse in the wilted

leaf and the bulliform cells show varying degrees of

buckling. Shields found that lateral and vertical

contraction in the bulliform and associated colourless

cells was no greater than in the adjacent rnesophyll.

In wilted leaves the abaxial ribs appear more pro-

minent as a result of the smaller amount of shrinkage

in the vascular tissue than in the surrounding cells.

Shrinkage of the bulliform and colourless cells appears

to play no part in involution except to facilitate the

turning inward of the upper leaf surface.

The fact that the bulliform cell walls are hygro-

scopic and, therefore, may assist in movement was

reported by Goossens & Theron (1934). This was

supposedly shown by the fact that dead, detached

leaf segments roll more tightly than in normal

reversible involution and, conversely, increasing their

water content leads to the partial unfolding of the

dead leaf segment. Shields (1951) comments that this

partial expansion of a dried leaf through adsorption

by all the cell walls (which are also dead in the

living blade) suggests that decreased hydration of the

cell wall may play a passive part in reversible wilting.

Since the bulliform cells are dead in this case,

absorbtion of water by the protoplasm and large

central vacuole cannot occur.

Involution does not result from plasmolysis.

Plasmolysis, through vertical contraction of the

rnesophyll, may cause an increase in width, probably

by relieving tissue strains through shrinkage of the

protoplasts (Burstrom, 1942). As a result vascular

bundles become more widely separated, and the

thick-walled, stiff abaxial epidermis, which does not

change in width, tends to curve backwards in plas-

molysis so that the underside of the leaf becomes

concave. Since plasmolysis involves turgor change the

diametrically opposed characteristic of inrolling in

wilted leaves implies that involution does not

result from turgor changes.

Loss of water from the bulliform cells to the

chlorenchyma in particular, may further contribute

to the collapse of these cells. With a cuticle on their

outer surface these epidermal cells are more resistant

than the rnesophyll cells to the drying effects of the

atmosphere. However, Haberlandt (1928) showed

that when an organ is transpiring rapidly, the

epidermis loses water to photosynthetic tissue with its

higher osmotic pressure.

Shields (1951) examined the bulliform cells in

unwilted but tightly rolled leaf buds. They are small

but turgid, rather than flaccid and collapsed, as in the

wilted blades of mature leaves. Furthermore, un-

folding from the developing bud involves general

growth, particularly in the adaxial rnesophyll, and

does not result primarily from the enlargement of the

bulliform cells, although these enlarge by stretching

growth due to turgor changes. The rolling mechanism

is, therefore, not related to the unfolding of the bud.

Rolling involves cohesion and shrinkage in both

living and non-living parts, and bud unfolding, turgor

and stretching movements at a time when all cell

walls are highly elastic.

To summarize, it has been shown that structural

elements other than bulliform cells contribute to

involution. 1. Rolling in drying is characterictic of

certain grass leaves entirely lacking in bulliform

cells, and if bulliform cells are present they may show

no buckling or other distortion in the wilted leaf.

2. Since average measurable lateral shrinkage (which is

negligible on the abaxial surface of wilted leaves)

amounts to 7-12 per cent on the adaxial rib face

(Shields, 1951) in naturally wilted leaves, subepidermal

sclerenchyma and other adaxial elements of the

rnesophyll must contribute to involution. 3. Since

dried leaves unfold partially when placed in water,

involution must result in part from decreased water

content in non-living cell walls which may or may not

contain protoplasts. Rolling cannot result entirely

from turgor movement, which, by definition, is a

reversible change in the water content of living cells.

Water loss by protoplasts alone is insufficient to cause

involution in a leaf composed largely of non-living

mechanical tissue. The form of the wilted leaf is

determined by a number of elements in the rnesophyll,

the buckling of bulliform and colourless cells, where

it occurs, being in part from passive compression.

Anatomical variations in the structure and distri-

bution of bulliform cells in different genera exhibiting

involutionary movements preclude the universal

application of any one explanation of this pheno-

menon in wilting grass leaf blades. What is important

from a descriptive point of view is the fact that the

outline of the lamina can be markedly altered by this

phenomenon. It appears to be constant within a

species and, therefore, it is not the degree of infolding

or unrolling that is important but rather the type of

movement.

This is confirmed by the work of Dunlop (1913) on

the curling of the leaves of different varieties of

sugar cane. Involute curling, where the upper surface

is protected, is characterized by the following anato-

mical structures: the bulliform cells are not prominent

and the cells immediately interior to these cells are

comparatively large and thin walled; the upper

epidermis is not greatly lignified. Varieties of sugar

cane exhibiting revolute curling, whereby the lower

surface is protected, have large prominent bulliform

cells attached to the vascular bundles by lignified cells.

The vascular bundles are closer together and the

upper epidermis is greatly lignified. Dunlop (1913) is

of the opinion that this revolute curling is a per-

manent characteristic of the varieties showing it.

A further possible function of the bulliform and

colourless cells has been advanced by Breakwell

(1915). He states that the arrangement and distri-

bution of the colourless cells allows light to penetrate

to the chlorenchyma cells, even in the rolled up

position of the leaf bud and thus aids in the develop-

ment of the leaf. The rapid development of leaves with

large bulliform groups and colourless parenchyma,

such as Astrebla pectinata, is cited as circumstantial

evidence for this possible function.

Epidermal cells in transverse section

Typical epidermal cells in transverse section

normally do not exhibit any particularly important

diagnostic characters, although examination of them

in section can be of assistance in the interpretation of

many epidermal structures as seen in surface view.

Sections through the epidermis can aid in establishing

the nature of the papillae, the attachment and

structure of other epidermal appendages, the

positioning of the stomata or the cuticle form and

thickness.

Epidermal cells are usually square to rectangular in

section, or the outer wall may be more or less arched

or even papillate. The papillate appearance, as seen

in surface view, may be due to a pronounced arching

of the whole outer tangential wall, or more commonly

(Vickery, 1935), due to the presence of a number of

separate papillae arranged in a longitudinal row or

rows. However, when these papillae are in a single

row, and as wide as the epidermal cell, the outer wall

108

A PROCEDURE FOR STANDARDIZING COMPARATIVE LEAF ANATOMY IN THE POACEAE.

I. THE LEAF-BLADE AS VIEWED IN TRANSVERSE SECTION

also appears as strongly arched in section. Narrower

papillae appear as distinct papillae on the outer

tangential wall surface. Where more than one row

of papillae are present on each cell they appear as

bifurcate or multiple papillae. The papillae may be

inflated and thin-walled, resemble conical warts or

may be sharp-pointed expansions of the outer wall

(Holm, 1891a) or teeth (Sabnis, 1921). It is also

common for the distal ends of the papillae to be

thickened.

The level of the stomata in different varieties of

sugar cane has been correlated with differences in

drought resistance by Merida (1970). They may be

located at the same level as the epidermal cells or

sunken below them as seen in transverse section. In

certain grasses, such as Spinifex hirsutus, the stomata

are present in depressions formed by the surrounding

cells being enlarged and raised above the general

level of the epidermal surface (Breakwell, 1915).

The presence of stomata on both surfaces, or only the

adaxial or abaxial surfaces can also be confirmed.

Micro-hairs are rarely seen in leaf sections but

prickles and macro-hairs are commonly sectioned.

The nature of the cushion cells surrounding the base

of many macro-hairs, whether raised or not, can

easily be determined. Interlocking prickles forming

arches over the stomata can sometimes be seen as in

certain Danthonia species (De Wet, 1960). Burbidge

(1946a) found hairs consisting of a bulbous-based

cell and a small apical cell imbedded in the epidermal

cells at the base of grooves of leaves of all species of

Triodia.

The size of the epidermal cells may vary over and

between successive bundles as well as the cells of the

adaxial and abaxial epidermides being of different

sizes. In many grasses, especially members of the

tribe Andropogoneae, the epidermal cells are excep-

tionally large, and can occupy up to half the leaf

thickness. These are included under bulliform cells.

Cuticle thickness can be determined in leaf sections.

Thus, it may be seen in many species that the lower

epidermis is more strongly cuticularized than the

upper. Slade (1970) has demonstrated the lack of a well

defined cuticle in shade-grown leaves of plants which

have distinct cuticles on the sun-grown leaves.

The thickness of this cuticle is of the utmost

importance in faecal analysis studies, because during

the digestion process all the cellulose cell-walls are

dissolved away leaving the cutinized cuticular mem-

brane behind. Thus, it is the relative thickness of the

cuticular membrane, and not of the entire outer

tangential wall, that is of importance in these studies of

the diet of grazing animals. It should be stressed

that the outer wall may be markedly thickened

but lack a thick cuticle. The cuticle proper, therefore,

consists of a layer of adcrusted cutin continuous

over the entire leaf surface. Each cell does not have an

individual cuticle but it may be individually thickened.

This difference can readily be determined in transverse

sections of these epidermal cells.

CONCLUSION

It is hoped that this attempt to introduce uniform

standards to the description of grass leaf blades, as

seen in transverse section, will stress the urgent need

for standardization in these studies. If this can be

achieved, anatomical descriptions will have a much

wider applicability in the fields of comparative leaf

anatomy and grass systematics in general.

ACKNOWLEDGEMENTS

The capable assistance of Miss L. Breytenbach and

Mrs J. Baker (nee Kimpton) with the line drawings

and the Division of Information, Department of

Agricultural Technical Services for composing the

keys is greatly appreciated.

UITTREKSEL

Omskrywende sleutels, definisies en diagramme, vir

die standaardisering, vereenvoudiging, en beskrywing

van die grasblaar struktuur soos beskou in dwarssnit,

word aangegee. Meer as 500 eienskappe is ingesluit

met die moontlikheid vir uitbreiding tot 999. Aan-

tekeninge van die variasie en die taksonomiese waarde

van die eienskappe word ook verstrek.

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Studies in the Hypoxidaceae. I. Vegetative morphology and anatomy t

M. F. THOMPSON* *

ABSTRACT

The leaf and corm morphology and anatomy of representatives of the genera Spibxene , Pauridia and

Empodium were studied. The corms are annual and tunicated, except in the group Aquaticae of Spiloxene. They

are swollen stems consisting of a number of internodes. In Spiloxene and Pauridia the roots grow from the base of

the corm, while in Empodium they develop from the sides. The epidermis of the older corms is replaced by several

layers of thin-walled cork. Characters of the corm coverings are used to divide Spiloxene into six groups. Four

leaf forms are recognised namely carinate, terete, canaliculate and plicate. Multicellular processes and unicellular

hairs occur occasionally.

The leaf stomata are paracytic. Most species have mucilage canals containing pectic compounds of mucopoly-

saccharides. The vascular bundles have complete or incomplete bundle sheaths and larger bundles have

sclerenchyma caps.

Resume

ETUDES SUR LES HYPOXIDACEAE. I. MORPHOLOGIE VEGETATIVE ET ANATOMIE

La morphologie et I'anatomie de la feuille et du bulbe des representants des genres Spiloxene, Pauridia et

Empodium ont ete etudiees. Les bulbes sont annuels et tuniques sauf dans le groupe Aquaticae de Spiloxene. Ils

ont des tiges gonfiees constitutes de nombreux internoeuds. Chez Spiloxene et Pauridia les racines sortent de

la base du bulbe , tandis que chez Empodium elles se developpent sur les cotes. L'epiderme des bulbes plus ages

est remplace par plusieurs fines couches de liege. Les caracteres de la couverture du bulbe sont utilises pour

diviser les Spiloxene en six groupes. Quatre formes de feuilles sont reconnues, a savoir : carinate , terete , cana-

liculate et plicate. Des processus multicellulaires et des poils unicellulaires se rencontrent occasionnellement .

Les stomates de la feuille sont paracytiques. La plupart des especes ont des catiaux mucilagineux conlenant des

composes pectiques de mucopolysaccharides. Les faisceaux vasculaires ont des etuis de faisceaux complets ou

incomplets et les faisceaux les plus grands ont des coiffes de sclerenchyme.

1 . INTRODUCTION

The Hypoxidaceae is represented in South Africa by

five genera, namely: Hypoxis L., Rhodohypoxis Nel,

Spiloxene Salisb., Empodium Salisb. and Pauridia

Harv. Empodium is sometimes included in the tropical

genus Curculigo Gaertn. and Spiloxene in Hypoxis.

Hypoxis and Rhodohypoxis occur mainly in the sum-

mer rainfall areas. Spiloxene and Pauridia are confined

to the winter rainfall region while Empodium is

widespread from the south-west Cape to the Transvaal

Drakensberg.

The genera are in need of taxonomic revision and

the aim of this study is to compare anatomical and

morphological features of the genera Spiloxene,

Empodium and Pauridia with special reference to

characters that could be of taxonomic value. This

paper covers the corm and leaf, while a second will

deal with the floral anatomy.

Previous anatomical work in the family was

largely confined to Hypoxis and Curculigo (Scharf,

1892; Schulze, 1893; Arber, 1925; Nel, 1914a;

Geerinck, 1968).

2. MATERIAL AND METHODS

Fresh material was collected in the field and voucher

specimens are housed in the Government Herbarium,

Stellenbosch (STE).

Fresh corms of Spiloxene capensis (L.) Garside, S.

aquatica (L.f.) Fourc., Empodium plicatum (Thunb.)

Garside and Pauridia minuta (L.f.) Dur. & Schinz were

cut longitudinally at different seasons to show the

development of the successive corms and the position

of the roots. The corms were cut at the beginning and

end of the growing season and during the dormant

period. Corms fixed in FAA were embedded in paraffin

wax, sectioned at 20-25 pm, stained with safranin

and fast green and mounted in Depex.

t Forms part of a M.Sc. thesis submitted to the University of

Stellenbosch (1972).

* Botanical Research Unit, P.O. Box 471, Stellenbosch.

The leaves of the following species were studied:

Spiloxene aquatica, S. capensis, S. flaccida (Nel)

Garside, S. minuta (L.) Fourc., S. ovata (L.f.) Garside,

S. schlechteri (Bol.) Garside, S. serrata (Thunb.)

Garside, Empodium plicatum, Pauridia longituba

M. F. Thompson and P. minuta. The Spiloxene species

were chosen to represent the groups defined by Nel

(1914a). The material used for the cross-sections of the

leaves was taken from the middle of the first two or

three foliage leaves (i.e. excluding the completely

sheathing leaves). Karpechenko’s modification of

Navashin’s fluid was found to be the best fixative. The

fixed material was embedded in paraffin wax, sectioned

at 15-20 pm, stained with safranin and fast green and

mounted in Depex.

3. THE CORM

(1) Morphology

(i) General

Two main types of underground storage organs are

found in the Hypoxidaceae of Southern Africa: an

erect, tuberous perennial rhizome in Hypoxis and

Rhodophypoxis and an annual corm in Spiloxene,

Empodium and Pauridia.

The annual corm is a swollen stem consisting of a

number of internodes formed by the apical meristem

of the shoot. There is no normal elongated stem and

the cataphylls, foliage leaves and axillary inflore-

scences arise directly from the upper half of the corm.

The branching is, therefore, monopodial and the

growth is usually orthotropic. In S. aquatica the

growth is usually somewhat plagiotropic i.e. horizontal

(Fig. 1 A), a condition brought about by the horizontal

growth of the apical bud. This differs from the

plagiotropic growth of the Iridaceae where the

branching is sympodial and a new corm is formed

from the uppermost axillary bud, the terminal bud

having produced a flowering shoot in the previous

season (Troll, 1937).

112

STUDIES IN THE HYPOXIDACEAE. I. VEGETATIVE MORPHOLOGY AND ANATOMY

together below by a ridge (Fig. IDa) and remaining

attached to the basal disc, branched or evenly latticed

at the base of the corm or reticulate, ending in a point

above (Fig. ID & IDa), e.g. S.flaccida.

(6) Pectinatae Ne1: Corm subovate, with basal discs,

fibres pectinate, not branched, apiculate, spreading at

the neck of the corm, normally adhering to the discs, if

separated from the disc then tending to remain joined

together at the base (Fig. IE), e.g. S. schlechteri.

(7) Minutae Nel: Corm basally flattened, without

basal disc, obconical, fibres adnate to the body of the

corm, not easily separable, producing a raised disc or

wide denticulate corona at the base (Fig. IF, lFa), e.g.

S. minuta.

With the exception of the Serratae, I regard these

groups as well-defined. The group Serratae, which Nel

separates from the Ovatae mainly because of its

serrate leaves, is not considered worthwhile upholding

as this character is variable in several of the groups. In

the Stellatae, for example, S. capensis has serrated i

leaves while the leaves of all the other representatives

are entire. The Serratae should thus be placed under

the Ovatae.

The description by Nel (1914a) of the structure of

the corm contains certain anomalies. He uses the term

Zwiebelknolle or bulbous corm and states that this

organ consists of leaf sheaths (p. 244 “besteht aus

Blattscheiden”). The corm is, in fact, a swollen stem

usually covered with the vascular remains of old leaf

sheaths. Nel also makes the erroneous statement that

the bases of the leaves may remain as discs (“Scheibe”)

Fig. 1. — Corm coverings in Spiloxene : A, S. aquatica (Aquati-

cae); B, S’. ovata (Ovatae); C, S. capensis (Stellatae);

Ca, detail of fibres of C; D, S'. ft accida (Flaccidae); Da,

detail of fibres of D; E, S. schlechteri (Pectinatae); Ea,

detail of fibres of E; F, S. minuta (Minutae); Fa, F seen

from below.

A new corm is formed annually on top of the old

one, the old shrunken storage organs of previous

seasons remaining at the base as discs (Fig. 2B). The

corm is either naked (in which case it is referred to as a

tuber) or it has a fibrous covering of tunics formed

from the remains of the sheathing foliage leaf bases

and cataphylls.

(ii) Spiloxene

As Nel (1914a, b) has shown, Spiloxene ( =Ianthe )

can be divided into groups on the basis of the fibrous

tunics of the corms. He recognized seven groups of

which six have tunicated corms and one a naked fleshy

tuber.

The groups of Nel (1914a, b) are given here with

modified diagnoses:

(1) Aquaticae Nel: Tuber fleshy, without basal discs,

never covered with fibres (Fig. 1A), e.g. S. aquatica.

(2) Ovatae Nel: Corm with discs, entirely covered

with branching fibres and twisting roots (Fig. IB), e.g.

S. ovata.

(3) Serratae Nel: Differs from the Ovatae in having

serrated leaves, e.g. S. serrata.

(4) Stellatae Nel: Corms with basal discs, fibres

easily separable from the old corm, much branched

below the middle or in the lower third and ending in

numerous pungent ribs pointing upwards (Fig. 1C &

ICa), e.g. S. capensis.

(5) Flaccidae Nel: Corm subovate with basal

discs, the upper (inner) fibres adnate to the corm, the

lower (outer) fibres easily separable but adpressed to

the body of the corm, spreading above, joined

Autumn Sprjng

Fig. 2. — Corms: showing the position of the roots in autumn

(A, C, E) and spring (B, D, F), after development of the new

corm. A-B, Spiloxene capensis (approx. xl,3); C-D,

Pauridia minuta (approx. x2,3); E-F, Empodium plicatum

(approx. xl,3).

M. F. THOMPSON

113

but this could possibly be a misprint for “Scheide”-

sheath. In his key and description of the groups Nel

describes all the corms except those of the Aquaticae,

as “discoideus”. This presumably refers to the old

corms which remain as discs at the base of the func-

tional corm. His description of the discs as two

lamellae with a nutrient rich space in between fits this

concept. In his description of the groups (1914b) he

does use the terms subdiscoid and subovate in

addition to discoid.

In autumn the new roots grow from the top of the

swollen corm, run under the tunics (where present) to

the widest part of the corm and pass through the

reticulations (Fig. 2A). After further development

during the winter growing season, it becomes clear

that these roots arise from the base of the present

season’s new corm. By spring the young corm has

become the main storage corm with the roots growing

from its base (Fig. 2B). The old corm remains as a

disc. In most Iridaceae, by contrast, the roots arise

from the base of a swollen storage corm and remain

functional the year after this corm has acted as a

storage corm (Troll, 1937; De Vos, 1970).

The position of the roots is the same in the autumn/

winter-flowering S. schlechteri and the spring-

flowering S. aquatica and S. capensis.

The old storage corm provides the nutrients for leaf

formation and, in the autumn flowering species, for

flowering.

An observation by Markotter (1936) that S.

schlechteri frequently reproduces vegetatively was

confirmed. In this species new corms appear to

develop from axillary buds on the base of corms, a

phenomenon seldom observed in other members of the

genus.

(iii) Empodium

E. plicatum has a corm with coverings resembling

those of the Stellatae group in Spiloxene, i.e. the

reticulate vascular fibres of the tunic are free at the

base. The dried-up intercostal tissue of the leaf bases

tends to remain longer than in Spiloxene, resulting in a

more scale-like covering, the reticulations being less

obvious.

The roots do not arise at the top of the summer

storage corm, but on its sides in the lower half and pass

immediately through the meshes of the tunics (Fig. 2E).

The roots develop after the first good autumn rains

and then the flowers appear before the leaves.

The new corm is formed by the apical meristem

above the old one in a monopodial orthotropic

fashion. Cormlets may be formed in axillary posi-

tions at the base of the developing corm towards the

end of the growing season (Fig. 2F).

(iv) Pauridia

Pauridia has a corm similar to that of the Flaccidae

group in Spiloxene, i.e. it is covered with reticulating

fibres which remain attached to the basal discs

formed by the previous seasons’ corms. As in Spiloxene

the roots arise at the base of the developing corm

(Fig. 2C).

(2) Anatomy

Anatomically the corms of the three genera,

including all the groups of Spiloxene, are very similar.

In the young corm the epidermis consists of a single

layer of thin-walled cells. Immediately interior to this

is a layer of parenchyma three to four cells deep which

lacks starch. The epidermis of the mature corm is

replaced by several layers of thin-walled cork cells

arising from a phellogen situated in the sub-epidermal

parenchyma (Fig. 3).

The main body of the corm is made up of a densely

packed ground tissue of parenchyma cells rich in

starch. Throughout the ground tissue there are

mucilage canals, which are not associated with the

vascular bundles. Canals are more numerous in the

mature corm than in the younger one. As in the leaf,

the canals are surrounded by radially flattened cells.

Scharf (1892) found that in Hypoxis the canals arise

schizogenously near the growing point and that they

have no membrane but are embedded in a ring of

smaller cells.

Numerous vascular bundles run irregularly through

the ground tissue and supply the roots and leaves. The

bundles normally have three to four tracheal elements

with ring or spiral thickenings.

Fig. 3. — Longitudinal section of the outer zone of the corm of S. capensis. c, cork; me, mucilage canal; r, root.

114

STUDIES IN THE HYPOXIDACEAE. I. VEGETATIVE MORPHOLOGY AND ANATOMY

4. THE LEAF

(1) General

The leaves in the Hypoxidaceae are radical with a

more or less sheathing base. The outermost leaves of

each year are reduced to bifacial leaves or cataphylls.

The foliage leaves are generally bifacial, but may be

unifacial and terete. In the bifacial forms the lip is

usually unifacial and represents the “Vorlauferspitze”

of Troll (1939).

The following description relates to the species listed

under Material and Methods and the main features are

summarized in Table 1.

(2) Anatomy

(i) Epidermis

The leaf epidermis of all the species studied consists

of a single layer of largely thin-walled cells, covered by

a smooth thin cuticle. In surface view the cells are

rectangular, elongated in the direction of the long

axis of the leaf, and have straight walls. The walls of

the epidermal cells along the leaf margin and keel are

sometimes thickened as in S. capensis (Fig. 4). Some

epidermal cells may be enlarged to form bulliform

cells as in the angle of the V of many of the carinate

leaf forms e.g. in S. serrata and P. minuta and in the

folds in Empodium. In most of the bifacial leaves the

cells on the adaxial surface are larger than those on

the abaxial side, but are not enlarged to form bulli-

form cells (e.g. S', ovata ).

In Spiloxene and Pauridia the stomata are scattered

on both surfaces, while in Empodium they are confined

to the intercostal zone. In all three genera they are

paracytic, i.e. with a single subsidiary cell parallel

to and flanking each of the guard cells (Fig. 5E). This

agrees with the observations of Stebbins & Kush

(1961), but Shah & Gopal (1970) found that in

Curculigo orchioides Gaertn. sixty per cent of the

stomata were tetracytic (i.e. with four to six subsidiary

cells). Shah & Gopal report further that in this species

the epidermal cells are not rectangular, as in the

hypoxid leaves which I have studied, but have an

irregular shape and arrangement.

The structure of the guard cells varies only slightly

in the species studied. The anterior cavity is well-

developed due to an outer ledge formed by the cell

wall and cuticle. The posterior cavity is poorly

Fig. 4. — Cross section of the leaf keel of S. capensis. me,

mucilage canal; ph, phloem; sc, sclerenchyma ; xy, xylem.

developed (e.g. S. aquatica ) or undeveloped (e.g.

S', flaccida and S. capensis). The guard cell walls are

unevenly thickened, as shown in cross-section (Fig.

5D). The median region of the wall bounding the

aperture and the entire opposite wall are thin and

elastic and together they constitute the hinges (vide

Eames & MacDaniels, 1947).

TABLE 1 . Comparison of leaf characters

(X) indicates partial or occasional presence. MC, mucilage canal; VB, vascular bundle.

M. F. THOMPSON

115

Fig. 5. — Leaf epidermis. A, epidermal processes on the keel of

S. capensis-, B, unicellular hair of E. plicatum ; C, multicel-

lular hair of S. capensis ; D, cross section of stoma of S.

flaccida; E, surface view of stoma of S. aquatica.

The serrations that occur along the margins of the

lower part of the leaf in S. serrata, and sometimes on

the margins and keel towards the tip of the leaf in S.

capensis, are multicellular epidermal processes.

In S. serrata the epidermis produces regular

incurved serrations two cells thick at the base. The

processes in S. capensis are very irregular. They

include the sub-epidermal thick-walled cells of the

margin and keel. They are often branched and may

extend into long pointed multicellular hairs (Fig. 5A

& C). Nel (1914a) reports that the teeth sometimes

contain chlorophyll but I found no chloroplasts in the

serrations.

On the ridges of the folds of Empodium plicatum

are unicellular hairs surrounded at their bases by

slightly raised epidermal cells (Fig. 5B) which do not

form part of the hair. These hairs resemble the cushion

macro-hairs of the Gramineae (Metcalfe, 1960).

The cuticle, thin throughout, continues over the

trichomes.

(ii) Mesophyll

The cells of the mesophyll are mostly irregular,

rounded to oval in shape, rich in chloroplasts and

with large inter-cellular spaces. Scattered cells contain

raphides and lack chlorophyll. The bifacial leaf forms

Df Spiloxene and Pauridia show a tendency to form

Dallisade tissue towards the adaxial side. There is no

Dallisade in the plicate leaf of Empodium. The cells on

;ither side of the midrib in the carinate leaves may be

mlarged and without chlorophyll as in S. minuta and

S', serrata. In the terete leaf of S. aquatica the meso-

ohyll has an aerenchymatous central part.

The mucilage in Spiloxene and Pauridia is confined

to mucilage canals which lie adaxially to, or, in

terete leaves centripetally to, the larger vascular

bundles (Figs. 4 & 6). The canals are bounded by

radially flattened cells. In Empodium the canals are

scattered in the mesophyll independently of the

vascular bundles. Mucilage canals are lacking in the

leaves of S. aquatica, although they are present in the

corm.

In sections of fresh leaves the mucilage showed no

reaction with Schultze’s solution and gave a reddish

colour with ruthenium red, indicating that the

mucilage is a pectic and not a cellulosic substance.

Tests with Alcian blue indicated the presence of

mucopolysaccharides. Scharf (1892) records that the

“colourless slime turns hard and brown when treated

with alcohol”. I found that the mucilage was dispersed

by fixatives containing alcohol and that it is soluble

in alcohol and not precipitated by it. Karpechenko’s

fluid (without alcohol) did not effect the position of

the mucilage.

(iii) Vascular bundles and sheaths

Cheadle & Uhl (1948) described five different types

of vascular bundles found in monocotyledons. The

larger vascular bundles of the species studied fall into

their Type IV which possesses one or two large central

metaxylem elements. These elements are tracheids

with ring or spiral thickenings. Cheadle (1968)

recorded primitive vessels with scalariform perfora-

tion plates in the roots of Hypoxidaceae but not in the

rest of the plant. The phloem consists of sieve tubes,

companion cells and phloem parenchyma, irregularly

arranged. The bundles often have sclerenchyma caps

at one or both poles (Fig. 4).

The smaller bundles generally lack sclerenchyma

and resemble Type I of Cheadle & Uhl (1948) which

has no larger metaxylem elements.

The vascular bundles are surrounded by a single-

layered bundle sheath of parenchymatous cells with

few or no chloroplasts. The sheaths may be (i)

complete, i.e. continuous round the bundle, e.g.

S. capensis (Fig. 4), or (ii) incomplete, i.e. interrupted

by sclerenchyma caps at one or both poles, e.g. in S.

flaccida and Empodium plicatum.

When the bundle sheath continues round the

mucilage canal to the adaxial epidermis, as in S.

capensis (Fig. 4), it may be regarded as a bundle sheath

extension.

(3) Leaf forms

On the basis of morphology and general anatomy

the leaves of the species studied may be divided into

four groups, namely, carinate, canaliculate, terete and

plicate (See Table 1).

(i) The carinate leaf

Examples: S. capensis, S. flaccida, S. minuta, S.

serrata, Pauridia longituba and P. minuta. (Fig. 6A,

6C).

The greater part of the length of the leaf is V-shaped

in cross-section with a large vascular bundle in the

angle of the V, forming a definite midrib and some-

times a marked keel as in S. capensis and S. flaccida.

S. serrata and S. minuta do not show as definite a keel

but the central bundle is still larger than the rest of the

bundles. In S. serrata and sometimes in S. minuta

the cells of the central adaxial mesophyll are enlarged

(up to 50 pm) and lack chlorophyll. The mucilage

canals lie adaxially to the larger vascular bundles,

except in S. minuta where they are lacking. The

bundles may have sclerenchyma caps as in S. capensis.

116

STUDIES IN THE HYPOXIDACEAE. I. VEGETATIVE MORPHOLOGY AND ANATOMY

Fig. 6. — Diagrammatic cross-sections of leaves demonstrating

leaf forms. A, Carinate: S. capensis, x 10; B, Terete:

S. schlechteri X 18; C, Carinate: S. serrata, X 18; D,

Canaliculate: S. ovata, x 10; E, Terete: S. aquatica,

X 10; F, Plicate: E. plicatum, X 10; me, mucilage canal;

ph, phloem; sc, sclerenchyma ; xy, xylem.

The sheathing part of the leaf does not have a

marked midrib. The tip of the leaf is a unifacial

“Vorlauferspitze” (Troll, 1939).

This is the most common type of leaf found in

Spi/oxene. Geerinck (1968), in comparing the leaves

of Haemodoraceae and Hypoxidaceae, regards this as

the typical hypoxidaceous leaf.

(ii) The canaliculate leaf

Example: S. ovata. (Fig. 6D).

The leaf is bifacial, U-shaped in cross-section and

without a midrib.

The adaxial epidermal cells are larger (about 50 /xm)

than the abaxial (about 30 yum). The mesophyll

consists of more or less even-sized irregular cells

without the formation of a palisade. Mucilage canals

bounded by radially flattened cells lie adaxially to the

vascular bundles. The 15-30 bundles are more or less

equal in size throughout the width of the leaf or the

alternating ones are somewhat larger. There is little or

no sclerenchyma associated with the vascular bundles.

This type of leaf is less common than the carinate

leaf and, among the species studied, only occurs in

S. ovata. Garside (1942) describes this type in S.

canaliculata Garside and considers the leaf shape

important in separating S. canaliculata from S.

capensis.

(iii) The terete leaf

Examples: S. aquatica and S. schlechteri (Fig. 6B, 6E).

The leaf is unifacial with a ring of ectophloic

bundles. The peripheral mesophyll is denser than the

central part which may be aerenchymatous (e.g. S.

aquatica). Mucilage canals, when present, lie centri-

petally to the vascular bundles.

There is a gradual transition from the bifacial,

canaliculate sheath to the unifacial terete part (Fig.

7a-d). The adaxial surface decreases gradually until

the entire leaf surface consists of the abaxial epidermis.

Fig. 7. — Transition from the bifacial leaf sheath to the unifacial

blade in S. aquatica. Cross-sections a-d taken at the points

marked on plant on left.

In the species investigated the terete condition

appears to develop differently from that in the

Haemodoraceae as described by Geerinck (1968)

where the leaves are “folded-welded”. Geerinck

described the Haemodoraceae as having a folded

(V-shaped in cross-section) basal part which fuses to

form a vertically flattened part with vascular bundles

alternating in direction, or a terete portion with

bundles in a ring with a peripheral phloem.

(iv) The plicate leaf

Example: Empodium plicatum. (Fig. 6F).

The leaf is bifacial and is folded longitudinally with

two or three folds on either side of the midrib.

Within the folds (ad- and abaxial) the epidermal cells

are enlarged to form bulliform cells. On the ridges the

the cell wails, especially the outer, are thickened.

Trichomes occur along the ridges and margins (Fig.

5B). Stomata are frequent in the inter-costal zones

of both surfaces. The mesophyll is uniform without

differentiation of a palisade. Mucilage canals occur

irregularly in the mesophyll and are not associated

with the vascular bundles as in Spiloxene.

Unlike the palm leaf, which has a prominent rib at

each fold (Tomlinson, 1961), not every fold has a

vascular bundle at the ridge. The midrib is the only

M. F. THOMPSON

117

large bundle in an abaxial ridge. The adaxial ridges

all have large bundles. Arber (1922) said that the folds

in Curculigo recurvata develop by alternating dorsal

(i.e. abaxial) and ventral (i.e. adaxial) invaginations of

the petiole or leaf sheath. The ventral sinuses occur

between the lateral bundles and opposite the midrib

while the dorsal sinuses occur opposite the main

lateral bundles (cf Fig. 6F).

5. CONCLUSIONS

The similarity in the vegetative anatomy and

morphology of Pauridia and Spiloxene suggests that

these genera are closely related. Pauridia has a corm

like the group Flaccidae of Spiloxene and there are no

major differences in the leaf structure of the two

genera.

The differences between Empodium and the other

two genera are greater. The corm, although super-

ficially similar, differs in the position of the roots. The

plicate leaf form is only found in Empodium. The

mucilage canals of the leaves are scattered in the

mesophyll in Empodium , while in Spiloxene and

Pauridia, where present, they are associated with the

vascular bundles (Table 1). The stomata are scattered

on both leaf surfaces in Spiloxene and Pauridia while

in Empodium they are confined to the intercostal zones.

Empodium has unicellular hairs while the epidermal

processes observed in Spiloxene, were all multicellular.

Within the genus Spiloxene the nature of the corm

coverings can be used to define groups (Nel, 1914a, b).

With the exception of the Serratae, which should be

combined with the Ovatae, the groups of Nel are

accepted.

The leaf form does not appear to be a good basis for

grouping species in any of the three genera, as other-

wise dissimilar species can have the same leaf form.

On the other hand, in the group Ovatae of Spiloxene

carinate [S', cuspidata (Nel) Garside, S’, gracilipes

(Schltr.) Garside], canaliculate [S', ovata, S. linearis

(Andr.) Garside] and terete [S. dielsiana (Nel)

Garside] forms occur. However, the type of leaf can

be used to distinguish between some closely related

species such as S', capensis and S. canaliculata (Garside

1942).

Apart from the fact that the internal structure

reflects the outward morphology, the leaf anatomy

was not found to provide characters of taxonomic

value. The presence of aerenchyma in S. aquatica is

probably a habitat response. Mucilage canals are

absent from the leaves in species as different as S.

minuta and S. aquatica. In Empodium however, their

arrangement is characteristic and different from the

other species investigated. The bundle sheaths are

generally indistinct and the presence of sclerenchyma

is not constant.

The morphology of the corm was found to provide

the vegetative characters most useful to the taxono-

mist.

UITTREKSEL

Die blaar- en knolmorfologie en anatomie van ver-

teenwoordigers van die genera Spiloxene, Pauridia en

Empodium is bestudeer. Die knolle is gerok ( uitsluitend

die groep Aquaticae van Spiloxene), eenjarig en bestaan

uit ’ n verdikte stingel met meer as een lit. By Spiloxene

en Pauridia groei die wortels uit die basis van die knol,

terwyl hulle by Empodium by die kante van die knol

uitgroei. In die ou knol word die epidermis deur dun-

wandige kurk vervang. Kenmerke van die knolomhulsels

word gebruik om Spiloxene in ses groepe te verdeel.

Vier blaarvorme word erken: gekiel, tereet, gekanaliseer

en geplooi. Multisellulere uitsteeksels en unisellulere

hare kom soms voor. Die huidmondjies by die blare is

paracyties. Slymkanale, met slym wat uit pektienver-

bindings van polisakkariede bestaan, word by die

meeste soorte aangetref. Die vaatbundels het volledige

of onvolledige bundelskedes en die groter bundels het

skier enchiemmusse .

REFERENCES

Arber, A., 1922. On the nature of the “blade” in certain

monocotyledonous leaves. Ann. Bot. 36: 329-351.

Arber, A., 1925. Monocotyledons — a morphological study.

Cambridge University Press.

Cheadle, V. I., 1968. Vessels in Haemodorales. Phytomor-

phology 18: 413-420.

Cheadle, V. I. & Uhl, N. W., 1948. Types of vascular bundles

in monocotyledons. Am. J. Bot. 35: 486-496.

De Vos, M. P., 1970. Bydrae tot die morfologie en anatomie van

Romulea: 1. Die knol. J. S. Afr. Bot. 36: 215-228.

Eames, A. J. & Macdaniels, L. H., 1947. An introduction to

plant anatomy. Ed. 2. New York: McGraw-Hill.

Esau, K., 1965. Plant anatomy. New York: John Wiley.

Garside, S., 1942. Plantae novae Africanae: Spiloxene canali-

culata Garside (Hypoxidaceae). J. S. Afr. Bot. 8 : 249-252.

Geerinck, D., 1968. Considerations taxonomiques au sujet des

Haemodoraceae et des Hypoxidaceae (Monocotyledons).

Bull. Soc. R. Bot. Belg. 101: 265-278.

Markotter, E. I., 1936. Die lewensgeskiedenis van sekere

geslagte van die Amaryllidaceae. Ann. Univ. Stellenbosch

14A, 2: 1-84.

Metcalfe, C. R., I960. Anatomy of the Monocotyledons. I.

Gramineae. London: Oxford.

Nel, G. C., 1914a. Studien iiber die Amaryllidaceae-Hypoxi-

deae, unter besonderer Berucksichtigung der afrikanischen

Arten. Bot. Jahrb. 51 : 234-286.

Nel, G. C., 1914b. Die afrikanischen Arten der Amarylli-

daceae-Hypoxideae. Bot. Jahrb. 51: 287-340.

Scharf, W., 1892. Beitrage zur Anatomie der Hypoxideen und

einiger verwanter Pflanzen. Bot. Zbl. 52: 152-327.

Schulze, R., 1893. Beitrage zur vergleichenden Anatomie der

Liliaceen, Haemodoraceen, Hypoxidoideen und Vellozia-

ceen. Bot. Jahrb. 17: 295-394.

Shah, G. L. & Gopal, B. V., 1970. Structure and development

of stomata on the vegetative and floral organs of some

Amaryllidaceae. Ann. Bot. 34: 737-749.

Stebbins, G. L. & Kush, G. S., 1961. Variation in the organiza-

tion of the stomatal complex in the leaf epidermis of

monocotyledons and its bearing on their phylogeny. Am. J.

Bot. 48: 51-59.

Tomlinson, P. B., 1961. Anatomy of the monocotyledons. II.

Palmae. Oxford: Clarendon.

Troll, W., 1939. Vergleichende Morphologie der hoheren

Pflanzen. Vol. 1, part. 2. Nachdruck, 1967. Koenigstein-

Taunus: Otto Koeltz.

.

Bothalia 12, 1: 119-121 (1976)

Notes on Veronaea including V compacta sp. nov.

M. C. PAPENDORF*

ABSTRACT

The genus Veronaea is discussed and a new species, V. compacta Papendorf described.

Resume

NOTES SUR VERONAEA INCLUANT V. COMPACTA SP. NOV.

Le genre Veronaea est discute el une nouvelle espece V. compacta Papendorf est decrite.

The genus Veronaea was erected by Cifferi &

Montemartini (1958) with type species V. botryosa and

characterized as follows:

Dematiaceus, Didymosporeus, Macronemeus.

Mycelium brunneum, dense ramosum, ex hyphis

brunneis, parce irregulariterque septatis, compositum;

conidiophora reptantia vel sub-assurgentia, consuete

ramosa, brunnea, septata, apicaliter fertilia, sursum

leviter incrassata aut clavata; conidia ex sterigmata

oriunda, hyalina, elliptica vel mediocriter elongata,

transverse 1-septata, dense botryoidea disposita;

sterigmata minuta, typice in cochlam retorta, etiam

irregulariter disposita.

Subramanian & Lodha (1964) described Sympodina

(Type: S'. coprophila), a Hyphomycete producing

brown, two-celled, solitary, dry conidia at the tips and

successively at the sympodially produced growing

points of a simple or branched, brown, septate

conidiophore. It has since been established (Papendorf,

1969; Ellis, 1971; De Hoog & Von Arx, 1973) that

S. coprophila is identical with V. botryosa. Subrama-

nian & Lodha ( l.c .) found that Sympodina is similar to

Pleurophragmium in its conidium ontogeny, but

different in being didymosporous and not phragmos-

porous like the latter. Furthermore, that Sympodina

appears to be congeneric with the Cladosporium sp.

illustrated by Barron & Busch (1962, Fig. 20-22)

and distinct from Scolecobasidium where the conidia

are borne on definite denticles.

In a revision of Scolecobasidium and Pleurophrag-

mium De Hoog & Von Arx (l.c.) compared a number

of genera characterized by conidia borne on denticles

and flat scars found on short ampulliform or more or

less elongated conidiogenous cells. According to these

authors Scolecobasidium is typified by ampulliform

conidiogenous cells bearing a limited number (1-3) of

two- to many-celled Y- or T-shaped conidia apically

on cylindrical denticles. It closely resembles Arxiella

which is characterized by reniform conidia with

cornute ends.

As a result of this delimitation they found it

necessary to transfer a number of Scolecobasidium

species as well as Veronaea simplex (Papendorf, l.c.)

to a new genus Ochroconis on the grounds that they

all have more or less elongated conidiogenous cells

with cylindrical denticles bearing 1-4-celled conidia

apically and laterally and often forming a distinct

rachis. The conidia are typical in having rounded ends

with a conspicuous flat basal hilum. Ochroconis

comes close to Dactylaria but in the latter the conidia

are fusiform with gradually tapering ends and the

denticles short and flattened. De Hoog & Von Arx

(l.c.) considers Pleurophragmium to be congeneric

with Dactylaria. In Phaeoisaria the conidia are also

* Institute for Botanical Research, Potchefstroom University,

Potchefstroom.

more or less fusiform but not pigmented and mostly

1 -celled. Veronaea closely resembles Ochroconis, but

differs in bearing its conidia on flattened scars which

are often pigmented.

Veronaea compacta Papendorf, sp. nov.

Figure 1 ,2.

Coloniae in agaro malti tarde crescentes ad diam.

1,5 cm quattuordecim diebus apud 25 °C attingentes,

effusae, gossypinae, ad partem mediam subelevatae,

subgriseae ad subfusce griseas, retrorsum fusco-

griseae; exudatum absens. Hyphae partim immersae,

ramosae, septatae, pallide sive dilute olivaceum

brunneae, 1, 5-3,0 pm diam. Conidiophora lateralia

sive raro terminalia in hyphis, saepe latiora quam

hypha fulciens atque usque ad 4 pm diam., macrone-

matoidea, mononematoidea, simplicia vel ramosa,

unicellularia sive multicellularia cellis saepe inflatis et

ampullaceis sive doliformibus, pallide ad olivaceum

brunnea, rare 50 /im longitudine excedentia; regio

fertilis definita rare amplius quam 10 yum in longitu-

dinem. Cellae conidiogeneae integrae, terminales

vel aliquando intercalares, polyblasticae, sympodicae,

cylindraceae ad doliformes sive ampullaceas, cicatri-

catae inconspicuis planisque cicatribus. Conidia in

racemulis nascentia, solitaria, arida, acropleurogena,

elliptica ad ovoidea sive oblonga ad subcylindracea,

apicaliter rotundata atque ad hilum latum truncatum-

que basaliter attenuata, glabra, obscure colorata ad

pallide brunea, continua vel uno septo medio praedita,

rare biseptata, saepe ad septum colligata, (4) 5-9 x

2, 5-3, 5 (4) jum, quoad rationes longitudinis latitu-

dinisque 2: 1-3: 1.

Colonies on malt agar slow growing reaching a diam.

of 1 ,5 cm in 14 days at 25 °C, effuse, cottony, slightly

raised in the centre, light grey to pale brownish grey,

reverse dark grey, exudate absent. Hyphae partly

immersed, branched, septate, pale to light olivaceous

brown, 1, 5-3,0 pm diam. Conidiophores lateral or

occasionally terminal on hyphae, often wider than

supporting hypha and up to 4 pm diam., macronema-

tous, mononematous, simple or branched, one- to

many-celled with the cells often inflated and flask-

shaped or doliform, pale to olivaceous brown, rarely

exceeding 50 in length, fertile region limited and

seldom more than 10 pm long. Conidiogenous cells

integrated, terminal or occasionally intercalary, poly-

blastic, sympodial, cylindrical to doliform or flask-

shaped, cicatrized with scars inconspicuous and flat.

Conidia produced in small clusters, solitary, dry,

acropleurogenous, ellipsoidal to ovoid or oblong to

subcylindrical, rounded apically and tapering to a

wide truncate hilum basally, smooth, faintly coloured

to pale brown, continuous or with a single median

septum, rarely 2-septate, often constricted at the

septum, (4) 5-9x2, 5-3, 5 (4) pm, length/width ratio

2:1-3 :1.

120

NOTES ON VERONAEA INCLUDING V. COMP ACTA SP. NOV.

r

/ / i 3* ' ; ^

(/ — _< V

/ l

Fig. 1. — Conidia and conidio-

phores of Veronaea com-

pacta, x2000.

Fig. 2. — Conidia and conidio-

phores of Veronaea com-

pact a.

M.C. PAPENDORF

121

Isolated from soil, Kosi Bay, Republic of South,

Africa, April 1974, M. C. Papendorf. PREM 44958,

dried culture on 1,5% malt agar, National Herbarium

Pretoria, Holotype. Transfers of the holotype have

been deposited in the Potchefstroom University

Culture Collection (No. 1222) and in the Centraal-

bureau voor Schimmelcultures, Baarn, Netherlands.

In this study Veronaea compacta was compared with

various isolates of V. botryosa, i.e. live subcultures of

the types of Cifferi & Montemartini (C.B.S. 360.65),

Subramanian & Lodha (C.B.S. 254.57) and an isolate

from decomposed cellulose (Papendorf & Jooste,

1974; C.B.S. 474.71). It was established that V.

compacta differs from V. botryosa mainly in the

characters of the colonies, conidiophores and conidia.

On malt agar the growth rate of the colony is consider-

ably slower than in V. botryosa, the colony diameter

for the two species being 1 ,6 and 2,8 cm respectively

after 14 days at 25 °C. The conidiophores are not as

long and flexuous as in V. botryosa but usually short

and aften compactly branched. The cells of the main

axis and its branches are mostly relatively short and

often inflated and wider than the supporting hypha.

The terminal conidiiferous region is limited and

produces only limited numbers of conidia in loose,

terminal clusters and never forms a long, conspicuous,

rachis-like structure of up to 200 pm as in V. botryosa.

On the whole the conidia of V. compacta are shorter

than those of V. botryosa and very rarely over 9 pm

long. The length/width ratio is 2: 1-3:1 while in V.

botryosa it reaches 4:1 or even 5:1.

ACKNOWLEDGEMENTS

I am indebted to Dr W. J. Jooste for his kind

assistance in preparing the photomicrographs and

drawings and to the C.S.I.R. for financial support

received.

UITTREKSEL

Die genus Veronaea word bespreek en ’ n nuwe

species, V. compacta Papendorf beskryf.

REFERENCES

Barron, G. L. & Busch, L. V., 1962. Studies on the soil

Hyphomycete Scolecobasidium. Can. J. Bot. 40: 77-84.

Cifferi, R. & Montemartini, A., 1958. Sui generi Muchmoria

Sacc e Veronaea n. gen. Atti. 1st. Bot. Univ. Lab. Crittogam.

Pavia , Ser. 5, 15: 67-72.

De Hoog, G. S. & Von Arx, J. A., 1973. Revision of Scoleco-

basidium and Pleurophragmium. Kavaka 1 : 55-66.

Ellis, M. B., 1971. Dematiaceous Hyphomycetes. Common-

wealth Mycological Institute, Kew, England.

Papendorf, M. C., 1969. New South African soil fungi. Trans'

Br. Mycol. Soc. 52 (3): 483-489.

Papendorf, M. C. & Jooste, W. J., 1974. New and interesting

records of South African fungi, Part VIII. Bothalia 11:

211-215.

Subramanian, C. V. & Lodha, B. C., 1964. Four new copro-

philous Hyphomycetes. Antonie van Leeuwenhoek 30:

317-330.

.

Bothalia 12, 1 : 123-127 (1976)

The soil mycoflora of an Acacia karroo Community in the

Western Transvaal

M. C. PAPENDORF*

ABSTRACT

A survey of the composition and distribution of the soil mycoflora of an Acacia karroo Community in the

Potchefstroom area was undertaken. A total of 858 speculating cultures representing 76 genera and 144 species

was recovered from this soil. The majority belong in the Fungi Imperfecti and only a limited number of Zygo-

mycetes and Ascomycetes and no Oomycetes or Basidiomycetes were recorded. Members of the genera Penicil-

lium and Aspergillus were the most abundant. The greatest concentration of individuals and species occurred in

the surface layers and a rapid decrease in numbers was noticeable with increasing depth. The nature of this

flora suggests a close correlation with the natural plant cover and the existing ecological conditions.

Resume

LA MYCOFLORE DU SOL D'UNE COMMUNAUTE D’ACACIA KARROO DANS LE TRANSVAAL

OCCIDENTAL

Une etude de la composition et de la distribution de la mycoflore du sol d'une communaute ef'Acacia karroo

a ete entreprise dans la region de Potchefstroom. Un total de 858 cultures sporulantes representant 76 genres et 144

especes a ete ex trait de ce sol. La majorite appartient aux Fungi Imperfecti et un nombre l i mite seulement de

Zygomycetes et d' Ascomycetes ont ete observes mais pas d'Oomycetes ni de Basidiomycetes. Les membres des

genres Penicillium et Aspergillus sont les plus abondants. La plus grande concentration d'individus et d' especes

se trouve dans les couches de surface et une rapide diminution en nombre a ete observee avec /’ accroissement en

profondeur . La nature de cette flore suggere une correlation etroite avec la plante naturelle de couverture et les

conditions ecologiques existantes.

During the past decades various aspects of the soil

mycoflora have been extensively studied in many

parts of the world and a voluminous literature has

accumulated (Alexander, 1970; Barron, 1968;

Doeksen & Van der Drift, 1963; Parkinson & Waid,

1960; Gilman, 1959; Burges, 1958; Chesters, 1949;

Domsch & Gams, 1970, etc.). As a result a fairly clear

picture of this flora has emerged and in recent years

the suggestion that it is of a uniform and cosmopolitan

nature (Waksman, 1916) has been criticized and partly

rejected in favour of the view that the soil mycoflora is

usually closely correlated with certain ecological

factors including the natural plant cover (Eicker,

1974; Morrall & Vanterpool, 1968; Park, 1965;

Clark, 1965; Garrett, 1963; Orpurt & Curtis, 1957;

Christensen et al., 1952; Tresner et al., 1954).

In South Africa the fungus flora of soils under a

natural vegetational cover has so far received little

attention and because of the great diversity of climate,

soil type and vegetation a vast and interesting field

remains to be investigated. Eicker (1969, 1970, 1974)

published a comprehensive list of fungi inhabiting

two forest soils in Zululand and the soil of the open-

savanna of the Transvaal.

This paper is a report on a study which was under-

taken to determine the composition and distribution of

the soil fungus population of an Acacia karroo

Community in the Western Transvaal. The Acacia

karroo Community is typical of the savanna of the

more arid western portion of the huge inland plateau

of the southern sub-continent commonly referred to

as the High Veld. It is dominated by Acacia karroo

Hayne and in this so-called Thorn Veld the Acacia

trees appear singly or in groups and are variously

scattered leaving large open spaces covered mainly

by grass species, herbaceous plants and shrubs

of different sizes. The following are the more common

species found in the area studied:

Trees: Ziziphus mucronata Willd., Ehretia rigida

(Thunb.) Druce, Rhus pyroides Burch., Maytenus

heterophylla (Eckl. &. Zeyh.) N. Robson.

* Institute for Botanical Research, Potchefstroom University,

Potchefstroom.

Shrubs: Grewia flava DC., Asparagus suaveolens

(Burch.) Miers., Asparagus cooperi Bak. and A.

laricinus Burch.

Climbers: Clematis brachiata Thunb., C. oweniae

Harv., Galium horridum Thunb. and Trochomeria

macrocarpa Hook.f.

Herbs and grasses: Teucrium capense Thunb.,

Kalanchoe rotundifolia Harv., Delosperma herbeum

N.E. Br., Sida dregei Burtt Davy, Lantana salvifolia

Jacq., Solarium nigrum Linn., Lippia scaberrima Sond.,

Conyza podocephala DC., Ehrharta panicea Sm.,

Digitaria eriantha Steud. and Themeda triandra Forsk.

The community concerned is situated on the

northern boundary of the present campus of the

University of Potchefstroom. The annual precipita-

tion in this area is c. 600 mm and occurs mainly in the

form of rain of which more than 80% falls during the

months October to March. The soil temperature

15 cm below ground level varies and fluctuates

between an annual summer maximum of c. 23 °C and

a winter minimum of c. 16 °C. The soil is a typical

red-brown loam consisting of the weathered products

of the underlying diabase.

METHODS

In order to obtain a composite and representative

soil sample of the community 10 sampling sites

covering the entire area were randomly selected. To

expose the profile a trench measuring 90x60 cm was

excavated at each site and from its sides soil samples

were taken at preselected depths of 0-2,5 cm, 15 cm

and 30 cm. Samples were collected by means of a

tapered and sharpened metal tube which could be

driven into the sides of the trench. Two cores were

taken from each level of the ten sampling sites. These

were placed in separate sterile containers and even-

tually combined in a large glass container, thoroughly

mixed and homogenized and subsequently sieved.

From this composite sample 25 g portions were

removed for further analysis. Peptone-dextrose agar

containing rose bengal and streptomycin (Martin,

1950; Johnson et al., 1959) was used as isolation

medium.

124

THE SOIL MYCOFLORA OF AN ACACIA KARROO COMMUNITY IN THE WESTERN TRANSVAAL

The isolation methods employed were all standard

procedures used in soil mycological investigations

and included the dilution-plate (Waksman & Fred,

1922), soil-plate (Johnson & Manka, 1961; Warcup,

1950) and the soil washing (Gams & Domsch, 1967;

Parkinson & Williams, 1961) techniques. Since

factors such as the method and media employed in

this type of investigation obviously have certain

limitations and selective influences it is not claimed

that the list compiled from existing data is a complete

inventory of this mycoflora.

RESULTS

During this survey a total of 858 sporulating isolates

was obtained and preserved in pure culture. An

analysis of these isolates gave the following distribu-

tion:

The individual species are listed in Table 1 .

TABLE 1. Fungi recovered from soil of Acacia karroo Com-

munity

TABLE 1. Fungi recovered from soil of Acacia karroo Com-

munity

M. C. PAPENDORF

125

TABLE 1. Fungi recovered from soil of Acacia karroo Com-

munity

DISCUSSION

In the final collection (Table 1.) the Zygomycetes,

Ascomycetes and Fungi Imperfecti are represented by

7, 8 and 61 genera and 8, 12 and 124 species respec-

tively. No Basidiomycetes or Oomycetes were

recovered from this soil.

It is generally accepted that the isolation of Basidio-

mycetes requires the application of specialized

techniques and media (Warcup, 1957; Robbins, 1950)

and consequently only very few, if any, Basidiomycetes

are recorded in most catalogues listing soil fungi

(McLennan & Ducker, 1962; Eicker, 1969, 1974;

Miller et a!., 1957; Wohlrab & Tuveson, 1965).

The absence of members of the Oomycetes should

probably be ascribed to unfavourable environmental

conditions including low soil moisture and high

summer temperatures often experienced in this area.

Eicker (1969, 1974) was also not able to isolate any

Oomycetes from two soil types in Zululand and only

obtained a single species of Saprolegnia from an

open-savanna soil.

The Zygomycetes seem to be fairly well established

in this habitat and are represented by the following

species: Absidia cylindrospora, Circinella sp., Cun-

ninghamella echinulata, Gongronella butleri, Mortie-

rella alpina , Mucor circinelloides, Rhizopus arrhizus

and Rhizopus sp. With the exception of Rhizopus the

zygomycetous genera recovered are each represented

by only one species. The total number of isolates (37)

for this group indicates that this soil is relatively

sparsely populated by Zygomycetes. The most com-

mon genus is Absidia (13 isolates) followed by Rhizo-

pus, Gongronella and Cunninghamella with 9, 5 and 4

isolates respectively. Circinella is represented by 3,

Mortierella by 2 and Mucor by only a single isolate.

Investigators studying the mycoflora of different

soil types from various localities have found that the

zygomycetous population can vary considerably. On

the whole cool, moist forest soils rich in nutrients and

energy sources seem to support a much larger and

more varied flora than poor and arid, sandy soils.

Many workers who have studied microfungi of

soils of more or less arid, sandy areas have indicated a

general paucity of mucoraceous forms (Wohlrab &

Tuveson, 1965; Kuehn, 1960; Nicot, 1960), Paine

(1927) also found that mucors are less prevalent in

open pasture land than in forests. Eicker (1969, 1974)

and Yung & Stenton (1964) reported fairly large

numbers of Zygomycetes from various soils. Goche-

nauer & Whittingham (1967) and Gochenauer &

Backus (1967) investigating the microfungi of certain

mesic forest and alluvial soils with coarse-grained

texture and low humidity reported that the most

important features of the populations of these habitats

were the rareness of members of the Mucorales and

Moniliales, other than Penicillium, in the dry sandy

soil as contrasted with the abundance of these types

in the mesic soil. This phenomenon is ascribed by them

to the high surface temperatures, low moisture

content and the limited energy sources available.

They consider these habitats to be too rigorous for the

survival of the generally hyaline and delicate types

as opposed to the demafiaceous forms and many

Sphaeropsidales which were recovered more abun-

dantly. The results of this investigation conducted in a

more or less semi-arid region, indicate that the

zygomycetous flora is not particularly varied and

rather poorly represented in number of recordings.

This seems to agree with the general pattern for the

soils from many arid regions.

Only a relatively small number of Ascomycetes were

recorded from this habitat. Warcup (1951b) pointed

out that the more generally used techniques, such as

the soil dilution method, produce only a few Ascomy-

cetes while methods where the soil is partially sterilized

by heat results in the isolation of more and a greater

variety of forms. By means of the more conventional

methods applied during this survey an aggregate of 14

isolates representing 8 genera were obtained. Only

one specimen of each of the following species was

recorded: Auxarthron umbrinum, Chaetomium pachy-

podioides, C. robustum, C. baineri, Herpotrichia

striatispora, Microascus cinereus, Neocosmospora vasin-

fecta, Westerdykella sp., Thie/avia sepedonium and

Sordaria fimicola. Chaetomium olivaceum and Chaeto-

mium sp. were represented by two isolates each. Of

the 14 isolates 11 originated from the upper horizon

while 3 were recovered from deeper levels.

126

THE SOIL MYCOFLORA OF AN ACACIA KARROO COMMUNITY IN THE WESTERN TRANSVAAL

In conformity with the general pattern of the soil

fungal population in a wide range of localities (Bhatt,

1970; Bagga, 1970; Eicker, 1969, 1974; Wohlrab &

Tuveson, 1965; Farrow, 1954) the Fungi Imperfecti

was found to be the largest and most varied group. A

total of 807 isolates representing 61 genera and 124

species was recovered from this soil. Most of the

genera were represented by a relatively small number

of isolates suggesting a limited occurrence in this

habitat. With only rare exceptions (Lim, 1969)

investigators reporting on the Fungi Imperfecti in

soil have found that Penicillium and Aspergillus spp.

are usually relatively abundant (Moubasher &

Moustafa, 1970; Eicker, 1969, 1974; Gochenauer &

Whittingham, 1967 ; Chen & Griffin, 1966; Christensen

et al., 1952.). In the Acacia karroo community

Penicillium was represented by 299 isolates including

25 species and Aspergillus by 69 isolates covering 7

species. Eicker (1969, 1974) also found that aspergilli

are less plentiful than the penicillia. Most abundant

were P. paraherquei (89 isolates), P. lilacinum (84),

P. adametzi (44), P. spinulosum (24), Aspergillus

ochraceus (50) and A. puniceus (10). Other genera

which seem to flourish in this habitat are Fusarium,

Humicola, Myrothecium, Trichoderma, Hyalotiella,

Dothichiza , Phoma and Beltrania.

Regarding the vertical distribution of soil inhabiting

fungi the available evidence (Eicker, 1970; Yung &

Stenton, 1964; Warcup, 1951a; Waksman, 1944;

Burges, 1958; etc.) indicates a general decrease in both

number of individuals and species with increasing

depth. In some reported cases particular fungi are

inclined to colonize certain horizons more readily

than others. Warcup (1951a) found that forms like

Gymnoascus, Cylindrocarpon radicicola and Paecilo-

myces were more abundant in the lower regions of

horizon A and seldom occurred near the soil surface.

On the whole the vertical distribution of the fungi

in the Acacia soil follows this pattern. The surface

layer produced 676 isolates against 158 and 24 for the

15 cm and 30 cm layers respectively. Only two species

were convincingly more abundant in the deeper

horizons, i.e. Penicillium adametzi and P. spinulosum,

while 19 were confined to these sub-surface layers.

From a taxonomic study of the various isolates it

was possible to erect 6 new genera and establish 11

new species (Papendorf & Von Arx, 1966; Papendorf,

1967a, 1967b, 1969a, 1969b; Papendorf & Du Toit,

1967; Van der Aa, 1967; Papendorf & Upadhyay,

1969; Stolk, 1969.). The presence of this relatively

large number of previously unknown fungi in this

particular habitat seems to support the view, referred

to earlier, that the soil mycoflora is often closely

correlated with certain ecological factors and the

natural vegetation.

ACKNOWLEDGEMENTS

My sincere appreciation is due to Dr J. A. von Arx

and members of the Centraalbureau voor Schim-

melcultures, Baarn, Netherlands, for the valuable

assistance received especially during my visits to that

Institute, and also to Drs G. C. A. van der Westhuizen

and W. F. O. Marasas of the Plant Protection

Research Institute, Pretoria, for their enthusiastic

co-operation. The financial support received from the

C.S.I.R. and the Department of Agricultural Technical

Services is gratefully acknowledged.

UITTREKSEL

'n Ondersoek na die samestelling en verspreiding van

die grondmikoflora van 'n Acacia karroo-gemeenskap

in die Potchefstroom-gebied is onderneem. In totaal

is 858 sporulerende kulture, wat 76 genera en 144

spesies verteenwoordig, in hierdie grond aangetref

Die meerderheid behoort tot die Fungi Imperfecti en

slegs enkeles tot die Zygomycetes en die Ascomycetes

terwyl geen Oomycetes of Basidiomycetes aangeteken is

nie. Lede van die genera Penicillium en Aspergillus was

in die grootste getalle teenwoordig. Die hoogste konsen-

trasie van individue het in die oppervlakkige grondlae

voorgekom terwyl 'n snelle afname met toenemende

diepte merkbaar was. Die aard van hierdie flora dui op 'n

noue korrelasie met die natuurlike plantbedekking en

die heersende ekologiese toestande.

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Soil Sci. 109: 247-249.

Barron, G. L., 1968. The genera of Hyphomycetes from soil.

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The soil microfungi of wet-mesic forests in Southern

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Stuttgart: Gustav Fischer.

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Garrett, S. D., 1963. Soil fungi and soil fertility. Oxford:

Pergamon Press.

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Press.

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willow and cottonwood lowland communities in southern

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Mycologia 59: 893-901.

Gochenauer, S. E. & Whittingham, W. F., 1967. Mycoecology

of willow and cottonwood lowland communities in southern

Wisconsin. I. Soil microfungi in the willow-cottonwood

forests. Mycopathol. Mycol. Appl. 33: 125-139.

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31: 123-158.

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M. C. PAPENDORF

127

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Botha Ha 12, 1 : 129-131 (1976)

The mycoflora of wheat field debris, Part II

W. J. JOOSTE*

ABSTRACT

Descriptions are given of four fungus species isolated from stubble of a wheat field in the Heilbron district:

Ascochyta hordei Hara, Ascochvta sorghi Sacc., Phoma pomorum Thiim. and Alternaria tenuissima (Fr.) Wiltshire.

Resume

LA MYCOFLORE DES DEBRIS D'UN CHAMP DE BLt

On decrit quatre especes de fungi isoles du chaume d'um champ de ble dans le district d' Heilbron. Ce sont :

Ascochyta hordei Hara , Ascochyta sorghi Sacc., Phoma pomorum Thiim. et Alternaria tenuissima {Fr.) Wiltshire.

The object of this study has been given in the first

part of this series (Papendorf & Jooste, 1974). The

four species described here have been deposited as live

cultures in the Potchefstroom University culture

collection and as dried down cultures in the Mycolo-

gical Herbarium (PREM) of the Plant Protection

Research Institute at 590 Vermeulen Street, Pretoria.

Materials and methods are similar to those

described previously. When required, cultures were

incubated under near U.V. light (330-480 nm) for

12h per day at 20 °C. The distance from the light

source to the cultures was 400 mm.

Ascochyta hordei Hara, in Sprague & Johnson,

Mycologia 42, 544-546 (1950); Sprague in Diseases of

Cereals and Grasses in North America, New York:

The Ronald Press Co., p. 156-158 (1950).

Figures 1, 2, 3 & 9

Colonies on 1,5% Malt extract agar growing

moderately fast, reaching a diameter of 80 mm in 12

days at 25 °C. Mycelium olivaceous-black to oliva-

ceous-grey, raised, floccose. Hyphae light to dark

olivaceous, septate, smooth or verruculose, varying in

diameter from 2-12,5 /tm, wide hyphae often thick-

walled and generally dark olivaceous, constrictions at

the septa common, branches often arising at right

angles. Pycnidia superficial, single or in clusters,

globose, subglobose, oblong-ellipsoidal or pyriform,

with a single neck and ostiole when young but

developing multiple necks and ostioles when older,

thin-walled, olivaceous to olivaceous-brown, hyphal

outgrowths on pycnidia 3-6 //m wide and up to 30 pm

long, young pycnidia with single ostioles up to 250 /mi

wide and 375 pm long with the neck up to 60 //m

long and 50 //m wide, extremely variable in dimen-

sions when older. Pycnidiospores ellipsoidal, tapering

to rounded ends, mostly straight, occasionally curved

or twisted, smooth, hyaline to light honey, 1 -septate,

rarely 2-septate, cytoplasm granular, 17, 5-25, Ox

4, 5-6,0 /mi, ave. 21 ,4x5,5 //m.

Specimen examined: P.U. Culture Collection No.

1 1 89, PREM 44954 dried down culture on 1,5% malt

extract agar.

According to Sprague & Johnson (l.c.) the spores

of this species can vary between 17-29 /mi in length

and 5-9 pm in width. It follows that this isolate is well

within the range of variation already recorded. The

main distinguishing character of A. hordei is the

* Institute for Botanical Research, Potchefstroom University,

Potchefstroom.

predominance of spores which taper from the middle

to the ends. The variation observed in the shape of the

pycnidia as well as the proliferation of necks and

ostioles at maturity is a feature not recorded before.

Ascochyta sorghi Sacc., in Sylloge Fungorum III,

406, (1884); Sprague & Johnson in Mycologia 42,

530-536 (1950), lists 7 synonyms; Sprague in Diseases

of Cereals and Grasses in North America, New York:

The Ronald Press Co., p. 159-161 (1950).

Figures 4, 5 & 10.

Colonies on 1 , 5 % malt extract agar growing slowly,

reaching a diameter of 60 mm in 13 days at 20 °C.

Mycelium raised, floccose, off-white to light buff.

Pycnidia formed only under near UV light. Hyphae

smooth, septate, slightly constricted at the septa,

hyaline to light honey, straight or flexuous, branches

arising at right angles, 2-5 /mi diam. Pycnidia super-

ficial, covered by hyphae, light umber, thin-walled,

globose, subglobose or ellipsoid, variable in size,

60-225 pm diam., with one or more terminal as well as

lateral ostioles, necks inconspicuous with cells

surrounding ostioles slightly darker. Pycnidiospores

hyaline, subcylindrical with ends rounded or sub.

fusiform and one end often truncate, usually 1.

septate, occasionally 2-septate, rarely constricted at

the septa, 12, 0-25, 0x2, 0-4, 0//m, ave. 17,5-2,7/tm.

Specimen examined: P.U. Culture Collection No.,

1191, PREM 44955 dried down culture on 1,5%

malt extract agar.

The pycnidiospores of A. sorghi was described by

Saccardo (l.c.) as oblong ovoid. Following the study

of a large number of isolates Sprague & Johnson (l.c.)

characterized the spores as “fusoid to ovate fusoid or

subcylindric and often constricted at the septum”.

According to this description the present isolate is

more cylindric or subcylindric than is characteristic

of A. sorghi. Furthermore, the one truncate end has

not been noted before. At present it is convenient to

refer to this isolate as A. sorghi since it does not

seem warranted to describe it as a new species.

Phoma pomorum Thiim., in Saccardo, Sylloge

Fungorum III, 152, (1884); Boerema, Dorenbosch &

Van Kesteren in Persoonia 6, 172 (1971); Boerema &

Dorenbosch in Studies in Mycology 3, 36-37 (1973).

Phoma prunicola (Opiz.) Wollenw. & Hochapt. in Z. Para-

zitK.de. 8, 595 (1936); Boerema, Dorenbosch & Van Kesteren in

Persoonia 4, 59-63 (1965); Dorenbosch in Persoonia 6, 9

(1970); Morgan Jones in C.M.I. Descriptions of Pathogenic

Fungi and Bacteria, 135 (1971).

Phoma bismarkii Kidd & Beaumont in Trans. Br. mycol. Soc.

10, 104-105 (1924).

130

THE MYCOFLORA OF WHEAT FIELD DEBRIS, PART II

Figs. 1-7. — 1, 2 and 3, Ascochyta

hordei pycnidiospores, pycni-

dia and multiple necked

mature pycnidia; 4 and 5,

Ascochyta sorghi pycnidio-

spores and pycnidia; 6 and 7,

Phoma pomorum pycnidia

and chlamydospores.

Figure 6, 7, 8.

Colonies growing moderately fast on oatmeal agar

reaching a diameter of 70 mm in 8 days under near UV

light at 20 °C. Mycelium on oatmeal agar delicately

cottony, olivaceous grey, pycnidia abundant. Hyphae

hyaline to olivaceous brown, smooth or verruculose,

septate, branched, 3,75-5,0 //m diam. Pycnidia glo-

bose to subglobose, partly submerged, thin-walled,

ostiolate with one or more ostioles, light to dark

umber, variable in size, up to 250 pm diam, neck

inconspicuous. Chlamydospores globose to sub-

globose, mostly intercalary, single or in chains,

verruculose or verrucose, thick-walled, umber to dark

umber, mostly continuous, rarely 1-septate, 10-14 /tm

diam. Dictyochlamydospores verrucose, dark honey

when young, umber when mature, terminal or inter-

calary, a mucous sheath often enveloping the spore,

irregulary obovoid, ellipsoid or obclavate, transversely

as well as longitudinally septate with a variable

number of septa. Pycnidiospores hyaline, ovoid to

ellipsoid, often slightly curved, continuous, with one or

more guttules, 5, 5-9, Ox 1 ,8-3,0 pm, ave. 6, 8x2, 8

pm.

Specimen examined; P.U. Culture Collection No.

1212, PREM 44956 as a dried down culture on oat-

meal agar.

P. pomorum is a common fungus with a wide host

range and is well known as P. prunicola (Boerema &

Dorenbosch, l.c.). The most important diagnostic

character is the presence of two types of chlamydo-

spores. The present isolate corresponds well with

those previously described; however, it varies in

having verrucose dictyochlamydospores which are

often enveloped in a mucous sheath.

On 1,5% malt-extract agar the growth of the

colonies are rapid reaching a diameter of 85 mm in 8

days under near UV light at 20 °C. The mycelium is

delicate, floccose and olivaceous brown. Numerous

pycnidia develop partly submerged in the agar.

This fungus has been recorded in South Africa as

Phyllosticta prunicola (Opiz.) Sacc. on Prunus arme-

niaca L. [Doidge in Bothalia 6, 629 (1950)].

Alternaria tenuissima (Fr.) Wiltshire in Trans. Br.

mycol. Soc. 18, 157 (1933); Neergaard in Danish

Species of Alternaria and Stemphyllium, Kopenhagen:

Einar Munksgaard, p. 145 (1945); Joly in Encycl.

Mycol. 23, 123-132 (1964) gives 37 synonyms; Ellis

in Dematiaceous Hyphomycetes, Kew: Common-

wealth Mycological Institute, p. 477 (1971).

Macrosporium tenuissimum Fr. in Systema Mycologicum

3,374 (1832).

Figure 1 1 .

Colonies growing rapidly on oatmeal agar reaching

a diameter of 80 mm in 10 days at 25 °C. Mycelium

olivaceous black, velvety, no pigment diffusing into

the agar and no evidence of crystals in the agar.

Hyphae light honey, smooth, septate, branched,

3,0-6, 5 //m diam. Conidiophores macronematous,

mononematous, isabelline, smooth, septate, often

branched with branches of unequal length, often

geniculate with 1-4 geniculations or a single terminal

W. J. JOOSTE

131

Figs. 8-11. — 8, pycnidiospores of

Phoma pomorum ; 9, pycni-

diospores of Ascochyta hor-

dei\ 10, pycnidiospores of

Ascochyta sorghi', 11, conidia

of Alternaria tenuissima.

conidiogenous cell, 40-155 /im long, mostly 40-60 x

4-5 pm, conidiogenous cells terminal, integrated,

sympodial, scars flat and slightly sunken, usually one

per cell. Conidia isabelline to dark isabelline, obcla-

vate, obovoid or ellipsoidal, smooth, verruculose to

verrucose, septate, generally with transverse, longitu-

dinal and diagonal septa, however conidia with only

transverse septa common, slightly to mildly constricted

at the septa, with or without false beaks, beaks

usually with a swollen or geniculate terminal coni-

diogenous cell, beak length usually not exceeding two

thirds of the length of the conidia, catenate with up to

8 conidia per chain, unbeaked spores 24-38x 12,5-

19,0 /an, ave. 30,0x14,5 //m, beaked spores 24,0-

84,0x11,0-19,0 pm, ave. 51,5x14,0 pm.

Specimen examined'. P.U. Culture Collection No.

1233, PREM 44957 dried down culture on oatmeal

agar.

According to Wiltshire ( l.c .) A. tenuissima was

originally published by Nees in 1818 on Kunze’s

specimen as Helmisporium (Helminthosporium) tenuis-

simum. Furthermore, Persoon in Mycol. Eur. 1,18

(1822) also quoted the Kunze specimen as H. tenuis-

simum. Fries (l.c.) made H. tenuissimum synonymous

with Macrosporium tenuissimum Fr. According to

Wiltshire (l.c.) Fries probably based M. tenuissimum

on specimen No. 212 of the Dezmazieres exsiccata

because of the fact that the conidia of Kunze's speci-

men in the Fries Herbarium lacks beaks while those of

Dezmazieres’s specimen is beaked. He proposed that

the Dezmazieres specimen be chosen to represent A.

tenuissima. It therefore follows that the Kunze

specimen, which was published by Nees in 1818 and

Persoon in 1822, should not be used as the type. This

was subsequently supported by Neergaard (l.c.). Ellis

(l.c.) indicated the authors of A. tenuissima as “(Kunze

ex Pers.) Wiltshire”. In view of the foregoing this

should be regarded as incorrect and A. tenuissima

(Fr.) Wiltshire accepted.

The isolate described here corresponds well with

descriptions of Wiltshire (l.c.), Neergaard (l.c.), Joly

(l.c.) and Ellis (l.c.) although the conidia are somewhat

smaller. Considering the variation in spore dimensions

given by Joly (l.c.) it is appropriate to regard it as

A. tenuissima.

ACKNOWLEDGEMENTS

The financial support of the Department of Agricul-

tural Technical Services and the C.S.I.R. is gratefully

acknowledged. Prof. M. C. Papendorf is thanked for

his interest and reading of the manuscript. Dr H. A.

van der Aa of the Central Bureau voor Schimmel-

cultures Baarn, Netherlands, is thanked for suggesting

the identity of the Ascochyta and Phoma species.

UITTREKSEL

Beskrywings word gegee van vier swamspesies wa{

geisoleer is uit stoppels van n koringland in die Heil-

bronse distrik: Ascochyta hordei Hara, Ascochyta

sorghi Sacc., Phoma pomorum Thiim. en Alternaria

tenuissima (Fr.) Wiltshire.

REFERENCES

Papendorf, M. C. & Jooste, W. J., 1974. The mycoflora of

wheat field debris, Part I. Bothalia 11 : 207-210.

Bothalia 12, 1 : 133-135 (1976)

Hans-Joachim Schlieben , collector extraordinary

O. A. LEISTNERt

ABSTRACT

Hans-Joachim Eberhardt Schlieben was born in Germany in 1902. Between 1930 and 1935 he collected plants

in Tanzania and since 1955 in Southern Africa, on Madagascar, the Mascarenes, the Comoro Islands and the

Seychelles. Almost 400 of the approximately 13 000 numbers which he has collected have been described as new.

Resume

HANS-JOACHIM SCHLIEBEN, RECOLTEUR EXTRAORDINAIRE

Hans-Joachim Eberhardt Schlieben est ne en Allemagne en 1902. Entre 1930 et 1935 il a recolte des plantes

en Tanzanie et depths 1955 en Afrique australe, a Madagascar, aux Mascareignes, aux Comores et aux Seychelles.

Pres de 400 nombres parmi les 1 3 000 qu'il a ainsi recoltes ont ete dccrites comme nouvelles.

“Captivating travel accounts, lessons in geography

and biology and, not least, philately stirred in me, even

as a schoolboy the desire to visit distant lands and

their people, especially the old German colonies with

their lush primeval forests and sun-drenched steppes”.

Thus he wrote in 1938 in a booklet on his Mt Kiliman-

jaro experiences looking back over the travels of his

youth and the historical plant collecting expeditions in

East Africa.

Hans-Joachim Eberhardt Schlieben was born at

Waldheim, Saxony, Germany on 26 May 1902, the son

of a medical doctor. After obtaining a horticultural

diploma, he gained experience in practical gardening

in Holland, England, France and Algeria, which he

visited between 1923 and 1928. During the next two

years he studied at the Horticultural College at

Geisenheim/Rhein and obtained a further diploma. He

continued his travels in central and southern Europe

and the Mediterranean countries visiting many

botanical gardens and horticultural establishments.

His keen interest in photography and writing resulted

in numerous illustrated articles in journals devoted to

gardening. The royalties he received helped to

finance further travels.

In 1930 his childhood dreams came true (Fig. 2)

when he journeyed to Tanganyika Territory

(Tanzania) to collect plants for the Botanical Museum

Berlin-Dahlem which was then under the directorship

of Dr L. Diels. He also collected for several other

European herbaria and thus ten or even more dupli-

cates were gathered of the majority of his numbers.

In Tanzania he met up with his only brother who

had left for Africa in 1926 and who had later settled in

East Africa as a farmer. His only sister accompanied

him on his expeditions from 1930 to 1932. Schlieben

collected widely in Tanzania (Table 1) and included in

his journeys areas such as the Uluguru Mountains, the

Mahenge Plateau and the Lindi District which were

botanically virtually unknown. During 1933 and 1934

he lived on Mt Kilimanjaro for seven months and

collected almost 1 000 numbers. Here he was visited

by Peter Greenway who was working at the Agricul-

tural Research Station at Amani at the time. The

last months of his stay in Tanzania were spent in the

Lindi District in the south of the country. In August

1935 he returned to Germany via Rhodesia and South

Africa where he paid short visits to the Botanical

Gardens at Kirstenbosch and Stellenbosch.

* After this article had gone to press, news was received that

Schlieben died on 14th July 1975 in Essen, Western Germany.

t Botanical Research Institute, Department of Agricultural

Technical Services, Private Bag XI 01, Pretoria.

TABLE 1. Schlieben collections (1930-1935)

During his five years in Tanzania Schlieben collected

close on 7 000 numbers and more than 50 000

specimens. Most of his first set in the Berlin-Dahlem

Herbarium was destroyed during World War II.

Fortunately duplicates of most of his collections are

preserved in other herbaria such as Kew, British

Museum, Paris, Brussels, Geneva, Zurich, Madrid,

Lisbon and other institutes including fhe National

Herbarium, Pretoria. The most complete set of his

Tanzanian collections is housed at Kew. Until 1955 he

remained in Germany where he worked for several

years at the Forestry Institute at Hamburg/Reinbek.

Here he investigated germination and seedling

development of tropical trees and started a collection

of wooden utensils which he curated.

Throughout these years he continued anting

articles and giving illustrated lectures. In 1941 he

published a book on his travels in East Africa entitled

“Deutsch Ost-Afrika einmal ganz anders.” It consists

of 436 pages, is illustrated by 182 of his own photos

and three maps and was published by Neumann,

Berlin. Together with K. Helbig he wrote a book on

Australia based on the diary of Ludwig Leichhardt,

one of the first explorers of central Australia between

1844 and 1847. This work was published in 1959 by

Brockhaus, Leipzig under the title “Schicksal im

australischen Busch”. His experiences on Mt Ki lman-

iaro are related in a booklet “Im Banne des Kiliman-

jaro”, which appeared in 1938 through the Neuer

134

HANS-JOACHIM SCHLIEBEN, COLLECTOR EXTRAORDINARY

Buchverlag, Dresden. On 16 December 1943 he

married Klara Schulte, a school teacher and daughter

of an ophthalmologist. Three daughters were born

from this marriage.

But the cold northern latitudes held too few

attractions for him and in 1955 he emigrated with his

family to South Africa to join his brother who had, in

the meantime, settled on a farm in the Soutpansberg,

Transvaal. On 1 February 1956 he was appointed as

Technical Officer in the National Herbarium, Pretoria

and he held this position until 31 January 1973.

During these years he continued collecting in his

official capacity, mainly in the Transvaal, Karoo,

Bushmanland and South-West Africa, while his long

vacations were spent on collecting expeditions to

Madagascar, the Mascarenes, the Comoro Islands and

the Seychelles. In 1971 he set out on a sentimental

journey to his old hunting grounds in Tanzania. In the

best collectors’ tradition one of the main aims was to

rediscover Aloe schliebenii, which he had found in a

remote region of the Nguru Mountains in 1933 and

which had not yet been described. In Nairobi he

renewed acquaintances with Peter Greenway and in

the Uluguru Mountains again met some of his old

assistants and porters (Fig. 3), who recalled many of

their past experiences and some of whom still remem-

bered his Kisuaheli name “Bwana Maua” — Mr

Flower. Old friends of the plant kingdom were also

found again, including Aloe schliebenii (Fig. 4).

Schlieben was among the first philatelists specializ-

ing in stamps with botanical subjects. Several medals

won on South African and international exhibitions

bear witness to his rich collections and his neat

presentations. Philately and gardening occupy much

of his time at present and he pursues these hobbies

with great enthusiasm and dedication.

He has more than 170 publications to his credit,

mainly illustrated articles on plants, travels and

philately published in German horticultural journals.

To date, he has collected close on 13 000 numbers

and more than 70 000 specimens. Some 400 of his

collections have been described as new and almost

100 of these have been named in his honour. The

name of Hans-Joachim Schlieben will be remembered

as long as scientists study plants from the eastern

parts of Africa.

UITTREKSEL

Hans-Joachim Eberhardt Schlieben is in 1902 in

Duit stand gebore. Vanaf 1930 lot 1935 het hy in

Tanzanie plante versamel en sedert 1955 in Suider-

Afrika, op Madagaskar, die Maskarene, die Comoro-

eilande en die SeycheUe. Bykans 400 van die amper

13 000 nommers wat hy versamel het, is as nuut

beskryf.

O. A. LEISTNER

135

Fig. 2. — Collecting in the Iringa

District, 1931. Mr Schlieben

is holding a fruit of the

woody climber Entada scan-

dens. The porters each carry

four plant presses.

Fig. 3. — Reunion in the Uluguru

Mountains, 1971.

Fig. 4. — Rediscovery of Aloe

schliebenii in the type loca-

lity in the Nguru Moun-

tains, 1971.

Bothalia 12, I: 137-160 (1976)

A phytosociological classification of the Nylsvley Nature Reserve

B. J. COETZEE*, F. VAN DER MEULEN*, S. ZWANZIGER*, P. GONSALVES* AND P. J. WEISSER*

ABSTRACT

The vegetation of the Nylsvley Nature Reserve in the Transvaal Mixed Bushveld is classified hierarchically

by the Braun-Blanquet Method of vegetation survey. The vegetation is seasonal grassland and deciduous savanna

with four floristically distinct major groups of plant communities: (1) grasslands and broad-leaved savannas on

non calcareous sandy soils on elevated sandstone and felsite areas; (2) microphyllous thorn savannas on calca-

reous, clayey, bottomland alluvial soils and termitaria thickets; (3) grassland and thorn savanna on calcareous

self-mulching vertic soils; and (4) secondary communities on long abandoned native settlements and recently

ploughed land.

Seven primary communities with 12 community variations and 4 subvariations, and three secondary com-

munities are described on the basis of 216 releves. The survey was carried out at two levels of detail, an ecosystem

study area in the broad-leaved savanna being surveyed in more detail, floristically and structurally, than the rest

of the Reserve.

Resume

UNE CLASSIFICATION PHYTOSOCIOLOGIQUE DE LA RESERVE NATURELLE DE NYLSVLEY

Une classification hierarchique de la vegetattion de la Reserve naturelle de Nylsvley dans le" Bushveld" Mixte

dit Transvaal a etc obtenue au moyen d'un releve execute suivant la methode de Braun-Blanquet. La vegetation est

faite de prairies saisonnieres et de savane perdant ses feuilles Thiver avec quatre groupes majeurs floristiquement

distincts de communautes de plant es:

1 ) des prairies et des savanes a larges feuilles sur des sols sablonneux non calcareux sur sites de gres et

felsite s\

2) des savannes epineuses microphylles sur des sols calcareux, argileux, au sous-sol alluvionnaire et les

bosquets de termitieres ;

3) des prairies et une savane epineuse sur des sols vertics calcareux ou I'herbe seche recouvre le sol d'elle

me me ( self mulching)-,

4) des communautes secondaires sur des emplacements que des populations indigenes ont abandonnes depuis

longtemps et qui ont ete laboures recemment.

Sept communautes primaires avec 12 variations de communautes et quatre sous-variations ainsi que trois

communautes secondaires sont decrites sur la base de 216 releves. L' inspection fat menee a deux niveaux de details,

une region destinee a T etude de Vecosysteme dans la savane a larges feuilles ayant ete examinee de plus pres au

point de vue floristique et structure! que dans le reste de le Reserve.

CONTENTS

Introduction

The Study Area

Climate

Soils

History and land use

Survey Methods

Distribution of sampling quadrats

Quadrat size, shape and analysis

Synthesis and results

Plant Communities

1. Communities of the elevated sandstone and felsite regions ( Rhynchelytrum villosum — Schizachy-

rium jeffreysiiTree Savannas and Grasslands)

1 . 1 Eragrostis pollens — Burkea Tree Savanna

1.1.1 Eragrostis pollens — Dombeya rotundifolia Variation

A Eragrostis pollens — Pollichia campestris Sub-variation

B Transitional sub-variation

C Transitional sub-variation

D Transitional sub-variation

Page

138

141

142

143

144

146

148

149

Botanical Research Institute, Department of Agricultural Technical Services, Private Bag X101, Pretoria.

138

A PHYTOSOCIOLOGICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

1.1.2 Eragrostis pallens — Setaria perennis Variation 149

1.1.3 Eragrostis pallens — Trachypogon spicatus Variation 149

1.2 Barleria bremekampii — Diplorhynchus Savanna 150

1.2.1 Pseudolachnostylis — Diplorhynchus Variation 150

1.2.2 Enneapogon scoparius — Diplorhynchus Variation 150

1.2.3 Burkea africana — Diplorhynchus Variation 150

1 .3 Eragrostis racemosa — Schizachyrium jejfreysii Tree Savanna and Grassland 151

1.3.1 Eragrostis racemosa — Combretum apiculatum Tree Savanna 151

1 .3. 1 . 1 Rhus leptodietya — Combretum apiculatum Variation 151

1 .3. 1 .2 Cymbopogon plurinodis — Combretum apiculatum Variation 151

1.3.2 Eragrostis racemosa — Digitaria monodactyla Grassland 152

1 . 3 . 2 . 1 Eragrostis nindensis — Digitaria monodactyla Variation 152

1 . 3 . 2 . 2 Tristachya rehmannii — Digitaria monodactyla Variation 153

2. Communities of flatbottomlands, excluding the self-mulching, vertic soils, and of termitaria

(Panicum maximum — Acacia tortilis Tree Savannas and Termitaria Thickets) 153

2.1 Sporobolus ioclados — Acacia tortilis Tree Savanna 153

2.2 Euclea undulata — Acacia tortilis Tree Savanna and Termitaria Thickets 153

2.2.1 Pappea capensis — Acacia tortilis Variation 155

2.2.2 Acacia nilotica — Acacia tortilis Variation 155

3. Communities of self-mulching vertic soils ( Aristida bipartita — Setaria woodii Tree Savanna and

Grassland) 156

4. Communities of abandoned settlements 157

Physiognomic Synopsis and General Remarks 159

Acknowledgements 160

UlTTREKSEL 160

References 160

INTRODUCTION

The Nylsvley Nature Reserve covers 3 120 ha in

Acocks’s (1953) Mixed Bushveld Veld Type. The

Reserve comprises the farm Nylsvley 560 KR,

situated between 24° 36' and 24° 42' S latitude and

28° 40' and 28° 44' E longitude, on the Springbok

Flats, 10 km south of Naboomspruit (Fig. 1). A

savanna ecosystem project has been initiated on part

of the Reserve by the Terrestrial Biology Section of the

South African National Programme for Environmen-

tal Sciences. The purpose of this primary survey was to

characterize the ecosystem study area floristically, to

show its relationships to the vegetation in the rest of

the Reserve and to supply basic information for

range utilization by larger ungulates. The survey also

formed part of the Terrestrial Biology Section’s

programme for inventorizing plant communities in

permanently conserved areas at a semi-detailed level

(Edwards, 1974). The Braun-Blanquet method (West-

hofif & Van der Maarel, 1973; Werger, 1974a),

adopted by the Botanical Research Institute for

primary semi-detailed surveys, was used. The survey

was carried out at two levels of detail. The more

detailed survey covers the areas chosen for intensive

ecosystem studies and the more general survey covers

the rest of the Reserve.

THE STUDY AREA

The Nylsvley Nature Reserve is situated in an

extensive undulating to flat terrain between 1 080 m

and 1 140 m altitude (Fig. 1).

Prominent hills are Maroelakop (1 140 m) in the

south and Stemmerskop (1 090 m) further north.

The small and shallow Nyl River runs across the

Reserve from south-west to north-east in a strip of

flat bottomland that divides the study area into a

number of gently sloping elevations. The south-

eastern elevations are underlain by sandstone, grits

and conglomerate bands of the Waterberg System and

the north-eastern elevations by felsite of the epicrustal

phase of the Bushveld Igneous Complex. The area has

a moderately low summer rainfall with a dry mildly

frosty, winter period. Several soil series occur,

associated with differences in geology and topography.

The uplands have non-calcareous, relatively sandy

soils with broad-leaved deciduous savannas and

seasonal grasslands. The bottomlands have calcareous

clay soils with microphyllous deciduous thorn

savannas and seasonal grasslands. Termitaria with

thickets are found throughout the felsite elevations

and parts of the bottomlands (Fig. 2).

The Reserve was formerly used mainly for cattle

grazing, but the natural state of the vegetation has been

generally well conserved. Small parts of the bottom-

lands were ploughed and disturbed patches around

abandoned settlements are still present on the sand-

stone elevations.

Climate

The climate, according to Koppen’s System, is a hot,

dry steppe with a dry winter period, designated

BShw (Schulze, 1947).

Large diurnal variation in temperature during the

cold winter months was regarded by Galpin (1926)

as among the main limiting ecological factors of the

Springbok Fiats. As shown in Fig. 3, the cool period

of water deficit stretches from April to September.

N v I st ro om

B J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

139

0 1 2 3 4 5 km

LEGEND

C o n t o u rs : 1 5,2 m interval Boundaries. Farms

6 0,9 m i n t e r va I Nylsvley ___

Roads Railways ---- Rivers -

Fig. 1. — Topographic map of the Nylsvley Nature Reserve and environs.

B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

141

Fig. 3. — Climate diagram based on data from Mosdene

590/486 and Nylsvley 590/307 Weather Stations (Weather

Bureau, 1954 & 1965), drawn after Walter & Lieth (1960).

p = mean monthly rainfall in mm

T = mean monthly temperature in °C

d = dry season

w = wet season

o = mean monthly rainfall over 100 mm (scale reduced

to 1/10).

Fig. 4. — A. Mean maximum and minimum temperatures at

Mosdene (Weather Bureau, 1954).

M = mean monthly maximum

D = mean daily maximum

d = mean daily minimum,

m = mean monthly minimum

B. Mean monthly rainfall at Nylsvley (Weather Bureau,

1965.)

Temperature

Temperatures recorded over the 10 year period

1920-1929 at the Mosdene 590/486 weather station,

10 km north-east of Nylsvley, give an indication of

temperatures that may be expected on the Reserve

(Weather Bureau, 1954; Fig. 4).

The station is situated at 1 097 m altitude in a

slightly elevated part of similar terrain. Ground frost

g may be expected daily in June and July, when mean

e daily minimum Stevenson Screen temperatures, which

= are considered as being generally 3 °C higher than

t ground minimum temperatures (Schulze, 1965), are

« 1,3 °C and 1,0 °C respectively. Occasional ground

frost occurs from May until September, during which

period mean monthly minimum temperatures range

from — 0,2 to 3,5 °C. These marginal freezing

temperatures suggest differences in frequency and

severity of frost with slight differences in topography

as a result of cold air drainage. This suggestion is

confirmed by Mr E. A. Galpin’s observations on

Mosdene (pers. comm.).

At Nylsvley cold air would accumulate in the

northern felsite area, which is at the lower end of a

broad, shallow valley originating at the Waterberg

Mountains 7 km to the west; in a moderately sloping

valley of the sandstone area and on the bottomlands.

Wind

Prevailing winds are east in summer and west in

winter (Weather Bureau, 1960). Mr E. A. Galpin

(pers. comm.) has observed that trees rapidly shed their

leaves after a drying westerly wind followed by cold

winds from the south. This effect is modified by

prevailing soil moisture conditions.

Rainfall

Rainfall data are based on 40 years’ records from

the Nylsvley 590/307 weather station, situated on the

Reserve at 1 097 m altitude (Weather Bureau, 1965).

The average annual rainfall is 630 mm. In about three

quarters of recorded years the rainfall exceeded 85%

of the normal (i.e. of the average for 30 years). An

average number of 62 days per year are rainy and

85-90% of the normal annual rainfall falls from

October to March (Weather Bureau, 1957).

Soils*

The soils on Nylsvley may be grouped into five

broad categories:

(a) Litholitic soils

These soils, found on the felsite and sandstone

elevations, consist of unconsolidated material or

hard rock with a minimum of soil particles.

(b) Non-calcar eons, well drained sandy soils

Soils of this category also occur on the elevated

parts. The well drained soils on felsite are shallow

with an orthic A-horizon on unconsolidated material

or rock, and belong mainly to the Mispah Series.

Well drained sandstone soils may be shallow,

belonging to the Mispah Series, or deep with orthic

A-horizons underlain by apedal B-horizons. The

Hutton Form is a medium to coarse sandy soil with a

red B-horizon and occurs on the upper slopes. The

Clovelly Form has a yellow B-horizon and is found on

lower slopes. Several series are distinguished and

together form a toposequence. The sequence from

* Information on soils was provided by Prof. H. J. von M.

Harmse and Mr P. J. Botha of the University of Potchefstroom,

who did a soil survey on Nylsvley.

142

A PHYTOSOCIOLOG ICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

upper to lower slopes is: distrophic, mesotrophic

and eutrophic Hutton soils, followed by distrophic and

mesotrophic Clovelly soils.

(c) Non-calcareous, poorly drained sandy soils

Poorly drained sandy soils occur at the footslopes

of the elevated areas and in elevated depressions. They

are classified into the Wasbank, Longlands, Fernwood,

Avalon and Mispah Forms. Wasbank and Longlands

soils have perched gley horizons overlying mottled,

plinthic B-horizons. Fernwood soils consist of a

shallow orthic A-horizon on regie sand and Avalon

soils have an orthic A-horizon, a yellow apedal B-

horizon and a soft plinthic B-horizon with mottling.

Klipfontein soils (Mispah Form) are shallow and

underlain by a hard flintite bank.

(d) Calcareous alluvial soils

The lowlands have a variety of alluvial soils. High

clay content soils occur furthest from the Nyl River;

lighter soils have been deposited closer to the river

beds; and along the streams relatively sandy river

banks have formed. The heavy soils, with more than

55% clay, belong to the Shephardvale Series and

lighter soils, with 35-55% clay, are classified as

Lindlay Series. Both series belong to the Valsrivier

Form. The soils have orthic A-and strongly structured

B-horizons. Soils of the Limpopo Series (Oakleaf

Form), with a weak-structured B-horizon have

developed on the river banks.

(e) Calcareous, vet-tic black clay soils

These deep, blgck soils occur in the bottomland

south of the alluvium. A vertic or melanic A-horizon

is characteristic. Three soil series are recognized here.

The Arcadia Series (Arcadia Form) is the most

extensive, consisting of a thick, strongly vertic A-

horizon on unconsolidated material or rock. As a

result of alternate wetting and drying the soil surface

tends to form very fine blocky mulch or loose granular

structures. This is a pure vertic soil with gilgai micro-

relief. The Gelykvlakte Series (Arcadia Form) differs

in that the soil surface is not mulching but crusting,

forming little cracks. Soils of the Bonheim Series

(Bonheim Form) have a melanic A-horizon, with

moderate to strong structure, overlying a dark-brown

structured B-horizon.

History and land use

The following account of the history and land use of

Nylsvley was extracted from an unpublished report

by Grunow (1974). The information was supplied by

Mr G. Whitehouse, previous owner of the farm

Nylsvley.

Cattle grazing system

In 1945, when Mr Whitehouse started farming on

Nylsvley, there were approximately 700 head of

cattle. In recent years the number has been reduced to

approximately 500 animals, representing a stocking

rate of approximately 6 ha/head. The number of herds

varied from three to four.

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B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

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The number of grazing camps has been gradually

increased to 25, the main camps ranging in size from

200-250 ha, except for one that is approximately

400 ha. Fifteen smaller paddocks serve, for example,

for keeping bulls and as maternity camps. Water

supply for the cattle comes from the Nyl River and

from two boreholes, situated in the bottomland near

the sandstone elevations.

The cattle were rotated, within limits set by water

supplies and field conditions, as determined by rain,

so that the whole farm was regularly grazed. The

most palatable grass species were not overgrazed and

rest periods for the vegetation were provided.

Three weeks after the first good rains, usually in

October, the cattle were moved to the savannas and

grasslands of the felsite area. From there they went to

the savannas of the extensive sandstone area, where

they stayed from mid-January until mid-May. The

latter area was divided into four camps and was

grazed by two herds of cattle of 1 50 animals each. Two

of the four camps were used during January and

February and the remaining two camps during March

and April. Grazing commenced on different camps

each year. Sometimes, when a dry period was followed

by heavy rain and the poisonous plant Dichapetalum

cymosum grew rapidly in the savannas of the sand-

stone areas, the cattle had to be temporarily removed

from these areas.

During the dry period of May until October the

cattle stayed in the bottomland. The vegetation of the

vertisols was used for cattle grazing between December

and June.

Non-domestic animals

The number of antelope and other game on the

farm varies considerably due to seasonal migrations.

Approximate numbers and fluctuations are: 150-500

impala; 40-100 kudu, which enter the Reserve in

December and stay for the summer; 20 duiker; 20

steenbuck; 8 reedbuck; 50-100 warthogs; and 40-200

monkeys and baboons. With improvea veld conditions

as a result of camping and stock reduction, impala

have become an increasing nuisance. The effective

rest period for a camp was usually shortened by the

impala, which grazed the camp before the cattle or,

preferably, thereafter when the grass was short. From

February to October the impala are found mainly in

the two northern camps of the sandstone area. There-

after they go north and enter the felsite area across the

bottomlands. When the marshy parts of the bottom-

lands are too wet to cross they stay in the lowveld

savannas.

The game are taken as the approximate equivalent

of 80 head of cattle, which gives a combined total

stocking rate of 5,4 ha/head for Nylsvley.

Fire

The previous owner never burned the pastures

intentionally. Incidental fires occurred over the whole

farm at irregular intervals and it is estimated that

approximately half the sandstone area burned once in

five years. Most fires occurred in the far northern camp.

Agronomy

Approximately 50 ha of the vertisol bottomlands

were cultivated and used regularly for cutting hay

from fodder crops. The area was banked off about 20

years ago to prevent it from being flooded.

Native settlements

The disturbance of native settlements, abandoned

since 1925, is still clearly apparent in the elevated

sandstone areas. Trees must have been cleared by the

occupants to construct huts and kraals and to win

areas for cultivation. The settlements were sited on

upland areas, well away from the bottomlands,

probably because the uplands are warmer and have

fewer mosquitoes.

SURVEY METHODS

The Braun-Blanquet method of sampling and

synthesis as described by Westhoff & Van der Maarel

(1973) and Werger (1974a) was followed, except that

in some instances quadrats were sub-sampled.

Distribution of sampling quadrats

Quadrats were distributed to include as much of

the variety in the study area as possible and to be

representative of homogeneous stands of vegetation.

The strategies used to achieve this in the intensive

study area differed from those in the rest of the

Reserve.

Intensive survey

The numbers of the 108 quadrats laid out in the

intensive study area are shown in Tables 1 and 2.

The quadrats were placed on or near grid intersections

and are permanently marked. A grid system was used

to ensure an even distribution of quadrats because of

the high sampling intensity. The flat topography and

tall trees also render orientation difficult. Grid points

were spaced 125 m apart on 345° (magnetic) compass

lines in the denser savanna and 250 m apart in the

more open savanna. The lines were 250 m apart and

follow a gentle gradient from upper to lower slopes.

Extensive survey

To cover the variation and to obtain a reasonably

even distribution of quadrats over the rest of the

Reserve, sample points were chosen in the field as

sampling proceeded. The number and approximate

position of the 107 quadrats laid out are shown in

Fig. 5.

Quadrat size, shape and analysis

It was estimated that quadrats of approximately

50 m radius are necessary to represent the structural

composition of most stands adequately, particularly

in the tree savanna of the intensive study area, where

trees are widely spaced. However, such quadrats are

unnecessarily large for the herb layer, which is ade-

quately represented in 10x20 m quadrats. Woody

species were, therefore, recorded in larger quadrats

where not adequately represented in 10x20 m

quadrats. Quadrat sizes are shown in Tables 1-6. In

small stands of vegetation with irregular boundaries,

as on some termitaria, quadrats of varying sizes and

shapes were used.

In the intensive study area a structural analysis was

made in addition to the standard analysis. The height

ranges of different layers of individuals were recorded

separately for every woody species. Cover-abundance

was recorded for each layer.

The following scale, taken from Westhoff & Van der

Maarel (1973), was used to estimate cover-abundance:

r: one or few individuals

+ : occasional and less than 1 %* of total plot area

1 : abundant and with very low cover, or less

abundant but with higher cover; in any case less

than 5 % of total plot area

2**: very abundant and less than 5% cover, or

5-25 % cover of total plot area

* Given by Westhoff etal. (1973) as 5% but this seems to be a

printing error (cf. Werger, 1974a).

** Subdivisions of this class are not shown in Tables 1-6.

144

A PHYTOSOCIOLOGICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

2m: very abundant

2a: 5-12,5% cover, irrespective of number of

individuals

2b: 12,5-25% cover, irrespective of number of

individuals

3: 25-50% cover of total plot area, irrespective of

number of individuals

4: 50-75% cover of total plot area, irrespective of

number of individuals

5: 75-100% cover of total plot area, irrespective

of number of individuals.

The scale as given by Werger (1974a) defines more

clearly what was understood as Class 1 : “numerous

but covering less than 1 % of the quadrat area, or not

so abundant but covering 1-5% of the quadrat area”.

The definition of class “r” in Werger’s (1974a)

account suggests a stricter interpretation of this

class than that held in this survey. The numerous

“r” in Tables 1-6 may therefore be taken as “+’\

which would correspond more with the normal

practice for allocating

Synthesis and results

The results of the synthesis are shown in Tables 1-6.

Each column represents a quadrat and thus a stand of

vegetation. The rows represent species occurring in

these stands. Cover-abundance symbols for every

species in each quadrat are entered in the matrix.

Species with similar distribution patterns are grouped

together and differentiate groups of stands, which

differ in species composition. The relationships

between the groups of stands are shown by the groups

of differential species they have in common. On the

basis of these relationships, closely related groups of

stands are combined to form a hierarchy of larger

groups of stands. Each stand group typifies a plant

community at a particular level in the hierarchy of

plant communities. Thus, the total species composi-

tion of any stand of vegetation, the characteristic

species combination of any plant community and the

various floristic relationships between individual

stands and between communities are shown in the

tables (Werger, 1974b).

Table 1 shows the community to which the intensive

study area and some releves from the rest of the

Reserve belong. Other communities in the Reserve are

shown in Tables 3-6.

Table 2 gives a structural analysis of the most

important woody species in the intensive study area.

The releves are arranged in the same sequence as in

Table 1 and lines between groups of releves are in the

same positions. Releve Nos. 211, 210, 209, 207,

208, 171, 172, 169 and 170 are omitted because no

detailed structural analysis was made.

Cover is presented in height classes. Several height

classes may be combined to a height range on the

bases of their similar distribution patterns. Table 2

shows how particular height ranges of some species

can differentiate between communities. Two groups of

species are shown:

(a) Species with height ranges that differentiate; and

(b) species with height ranges that do not differentiate.

In many instances a single layer of individuals

recorded in the field extends across more than one

height class in Table 2. The cover-abundance value in

such a case is the total for the height classes concerned.

This is indicated in the table by entering the cover-

abundance value in the lowest height class and an

asterisk in the higher classes. For example, if a layer

of individuals was recorded in the field as Burkea

africana, 4-6 m, with cover-abundance of 1, this

value is entered as “1” in the row for B. africana,

4- 5 m, with an asterisk in the row for B. africana,

5- 6 m.

PLANT COMMUNITIES

The vegetation of the Nylsvley Nature Reserve is

floristically divided into four major groups of com-

munities:

(1) communities of the elevated sandstone and

felsite areas;

(2) communities of termite mounds and of the flat

bottomlands, excluding self-mulching, vertic soils;

(3) a bottomland community on self-mulching,

vertic soils; and

(4) communities of abandoned settlements.

The plant communities are mapped in Figure 6.

1. Communities of the elevated sandstone and felsite

areas (Tables 1-3) (Rhynchelytrum villosum —

Schizachyrium jeffreysii Tree Savannas and Grass-

lands)

The elevated sandstone and felsite regions have

litholitic and shallow to deep, non-calcareous sandy

soils. The vegetation comprises broad-leaved savannas

and grassland. Savannas occur on the upper parts,

where frost is relatively mild and soils are well

drained. Grasslands occur on frostier lower slopes

with impeded drainage. The differential species are

given in Tables 1 and 3. The most common ones are

the grasses Rhynchelytrum villosum and Digitaria cf.

eriantha*

The group of species which differentiates this

community is similar to the group that differentiates

the Loudetia simplex — Aristida aequiglumis Wood-

lands, Shrublands and Grasslands on leached soils in

the Rustenburg Nature Reserve (Coetzee, 1975),

where the annual rainfall is approximately 100 mm

higher than at Nylsvley. Related communities in

similar rainfall areas were also described by Du Plessis

(1972), Theron (1973) and Coetzee (1974). Marked

differences with the Nylsvley community include, for

example, the occurrence and high constancy of

Loudetia simplex at Rustenburg and the high presence

of Rhynchelytrum villosum and Schizachyrium jeffreysii

at Nylsvley.

An example of closely related vegetation in an

area where the rainfall is 100 mm lower than on

Nylsvley is provided by Rutherford (1972) from the

central portion of the Omuverume Plateau in South

West Africa. Half the group of differential species of

the Nylsvley Rhynchelytrum villosum — Schizachyrium

j

Braun Blanquet cover abundance values (see: Survey methods)

□ '

Key to Fig. 7.

* To be confirmed.

B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

145

Eragrostis pallens-Pollichia campestris Sub-variation (u.iAa)

Eragrostis pallens-Pollichia campestris S u b- v a r i a t i o n (u.lAb)

Eragrostis pallens-Trachypogon spicatus Variation (1.1.3)

O c h n a

p u I c h r a

Fig. 7. — Variations of Eragrostis pollens — Burkea Tree Savanna. Relative frequency of cover-

abundance classes in different height layers of dominant woody species. Full width of column

represents 100% frequency (simplified after Table 2). See key on opposite page.

146

A PHYTOSOCIOLOGICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

jeffreysii community and several more species in the

characteristic species combination of the Eragrostis

pallens — Burkea Tree Savanna (see I . I) are included

in Rutherford’s table of vascular plant species for his

Burkea- Digit aria Woodland-savanna.

The Rhynchelytrum villosum — Schizachyrium jeffrey-

sii community has three sub-divisions associated with

clear topographic and soil differences. A group of

species, which is restricted to this community but is

seemingly sensitive to frost, and/or poor soil drainage,

cuts across the three sub-divisions. This species

group is absent from the lower slopes of the sandstone

and felsite elevations and includes: Phyllanthus

parvulus, Combretum t nolle , Euclea natalensis, Dom-

beya rotundifolia, Ozoroa paniculosa and Asparagus

suaveolens. Other species indicate reticulate relation-

ships between pairs of the three main groups.

1 . 1 Eragrostis pallens — Burkea Tree Savanna (Tables

1 and 2)

This community occurs on the non litholitic, sandy

soils in the higher parts of the sandstone area. Soils

are of the Mispah, Hutton and Clovelly Forms,

which are moderately shallow to deep, well-drained

and highly leached. The soils are mainly yellow-brown

to red and have a low nutrient status. The topography

is flat to very gently sloping, usually less than 2°.

Occasional sandstone outcrops occur, but the degree

of rockiness is always less than 1 % of the plot size.

Signs of shallow digging, small animal holes and

antbear burrows are common. Usually there is a

5-10% cover of plant litter, consisting of leaves,

mainly of Burkea africana and Ochna pulchra , grasses

and twigs. Occasional dead trees lie on the ground.

The number of vegetation layers, their height and

cover varies in the different community variations.

A structural analysis is given in Table 2 and Fig. 7.

The community is differentiated by a large group

of species as shown in Table 1. Constant differential

species in the tree and shrub layer are Grewia flave-

scens, Strychnos pungens, S. cocculoides, Lannea

discolor and Securidaca longipedunculata; and in the

field layer, the grasses Eragrostis pallens, Aristida

argentea and A. stipitata and the herbs Vernonia

poskeana , Limeum viscosum, C/eome maculata, C.

rubella and Dichapetalum cymosum.

Galpin (1926) described the Eragrostis pallens —

Burkea community as the Sandveld Association of the

Springbok Flats. The woody component of the

community corresponds very well, structurally and

floristically, with Galpin’s description. However,

Galpin stated that “No specific grass can be said to

dominate this Association”.

The community consists of three variations, one of

which has several sub-variations. The variations and

sub-variations are correlated with a gradient from

upper to lower slopes and with corresponding soil

and climatic differences.

1.1.1 Eragrostis pallens — Dombeya rotundifolia

Variation

This variation is the most extensive and occupies

approximately 250 ha on the upper and middle

slopes of the sandstone area, where the soils are well

drained. The variation occurs on soils of the Hutton

and Mispah Forms and on soils transitional to the

Clovelly Form. Total tree cover varies from 20-60%.

Trees range in height from seedlings and low juveniles

up to approximately 15 m. The dominant trees are

Burkea africana, Terminalia sericea and Combretum

mode and the dominant shrubs are Ochna pulchra and

Grewia flavescens. As shown in Table 2 and Fig. 9

the height ranges at which these species generally have

their maximum cover vary considerably between

species and between the different community sub-

variations. An average of 14 woody species was

recorded per plot. Grass and forb cover varies from

15-65%.

The community variation is differentiated from the

two other variations on less well-drained soils and

lower sites by Phyllanthus parvulus, Combretum molle,

Euclea natalensis, Dombeya rotundifolia, Ozoroa

paniculosa and Asparagus suaveolens. Elsewhere in the

Reserve these species also differentiate between

communities on well drained uplands and poorer

drained bottomlands (Table 3). This variation is also

structurally differentiated (Table 2) by some woody

species, which are taller here than in the other two

variations. These are: taller than 2 m Lannea discolor-,

taller than 2,5 m Ochna pulchra-, and taller than 8 m

Terminalia sericea. In the other two variations some of

these species occur as coppicing shrubs with emergent

dead twigs. As the twigs showed no sign of fire

damage it seems likely that this is caused by accumula-

tion of cold air and/or impeded soil drainage.

Three sub-variations, two of which are of limited

extent, occur.

1 . 1 . 1 A Eragrostis pallens — Pollichia campestris Sub-

variation

The largest and most typical part of the Eragrostis

pallens — Dombeya rotundifolia Variation occurs on

eu-, meso-, and distrophic Hutton soils of the middle

and upper slopes.

This sub-variation includes a primary and secondary

phase:

(a) The primary phase is characterized by the

regular occurrence, and gradual change in cover

values, of woody species over their entire height range

(Table 2; Figs. 7 and 8).

There is a marked low occurrence of woody species

in the 3-3,5 m height range where the tallest shrubs

are mixed with low trees. Below this height range

shrubs are dominant and above this height range trees

are dominant, with a regular pattern of alternating

dominant species:

Above 8 m: Burkea africana, followed by Termi-

nalia sericea.

3,5-8 m: Burkea africana, followed by Termi-

nalia sericea, Ochna pulchra and

Combretum mode.

3-3,5 m: Low trees of Terminalia sericea,

followed by Burkea africana and

Ochna pulchra.

2-3 m: Grewia flavescens and Ochna pulchra

(tall shrubs).

1-2 m: Ochna pulchra, followed by Grewia

flavescens.

Below 1 m : Ochna pulchra shrubs.

Burkea africana is the most common young tree in

the shrub layers below 3 m, followed by Terminalia

sericea and Combretum mode, which are most abun-

dant at 1-2 m and at 0-0,5 m. The field layer, which

has a total cover of 15-60%, is dominated by Era-

grostis pallens and Digitaria cf. eriantha.

The sub-variation is differentiated by a group of

species with low cover. Constant differential species

are, in the tree and shrub layer, Peltophorum africanum

and Mundulea sericea and in the field layer, Panicum

maximum, Pollichia campestris and Phyllanthus made-

raspatensis. The grass Panicum maximum clusters

under large trees and dense Grewia flavescens shrubs,

TABLE 1. Eragrostis pal lens — Burkea Tree Savanna.

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B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

1 47

Fig. 8. — Vegetation structure in

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Eragrostis pallens — Pollichia

campestris Sub-variation.

Fig. 9.— Vegetation structure in

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Eragrostis pallens — Pollichia

campestris Sub-variation.

148

A PHYTOSOCIOLOGICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

In addition to the floristic similarities with the

Burkea-Digitaria Woodland-savanna on the Omuve-

rume Plateau already mentioned, there are similarities

in structure and dominant woody species. The

dominant woody species given by Rutherford (1972)

are: Burkea africana ; Terminal ia sericea ; Combretum

psidioides subsp. dinteri; and Ochna pulchra, which is

mentioned as very common and usually less than 3 m

tall. In the Eragrostis pallens — Pollichia campestris

Sub-variation at Nylsvley Combretum molle replaces C.

psidioides and Grewia flavescens is also a very common

woody species less than 3 m high. The low mounds

occupied by Grewia flavescens at Nylsvley occur also at

Omuverume where they are occupied by Combretum

collinum. Both species generally form thickets on low

mounds.

The dominant grasses in the Nylsvley community

are Eragrostis pallens and Digitaria cf. eriantha.

On the Omuverume Plateau, the Burkea-Digitaria

community is considered to be in a near climax stage,

probably because of protection from large grazing

mammals by an encircling barrier of cliffs. The

dominant grasses there are Digitaria po/evansii,

Brachiaria nigropedata, Andropogon gyanus and A.

schirensis. On an adjoining plateau with seemingly

identical abiotic conditions, but with cattle grazing,

Rutherford (pers. comm.) noted that Eragrostis

pallens was the dominant grass, as on Nylsvley. The

other dominant grass on the grazed plateau was

Arstida stipitata, which at Nylsvley has a high

presence, occasionally with high cover. The high cover

of Eragrostis pallens at Nylsvley seems, therefore, to be

a result of grazing.

Minor structural units within the primary phase are

(Table 2):

(i) Areas where 1 ,5-3 m Grewia flavescens shrubs

cover 1-10% and where Euclea natalense and

Cassia mimosoides have high presences.

(ii) Areas where 1 , 5-3 m Grewia flavescens shrubs

cover less than 1 % and where Vitex rehmannii

has a high presence.

(b) The secondary phase occurs in small zones

around old settlements (Fig. 9).

This secondary vegetation is typified in Releve

Nos. 47, 55, 57 and 59. Soils here are eutrophic,

of the Hutton Form, and relatively rich in phosphate

content. Plant litter of Burkea africana leaves, twigs

and grasses covers up to 35% of the soil surface.

The vegetation structure differs from the primary

phase in that trees are less regularly distributed over

their entire height ranges; high cover values occur in

discrete layers; and Burkea africana and Terminalia

sericea both have their maximum cover values at

5-10 m. Total cover in the dense layer is between 20

and 55%. Dominant woody species at various heights

are as follows (see Fig. 7 and Table 2):

Above 5 m: Burkea africana , followed by Termi-

nalia sericea.

3 , 5-5 m : Terminalia sericea, followed by Burkea

africana.

3-3 ,5m: Terminalia sericea (tall shrubs), fol-

lowed by Combretum molle (low trees).

2-3 m: Grewia flavescens, followed by Termi-

nalia sericea and Ochna pulchra.

Below 2 m : Ochna pulchra.

Young trees of Burkea africana, Terminalia sericea

and Combretum mode occur in the shrub layers. Cover

in the field layer is between 5 and 50% and Panicum

maximum is the dominant grass, followed by Digitaria

cf. eriantha, Eragrostis pallens has less than 1 % cover.

Peltophorum africanum, Mundulea sericea and

Maytenus heterophylla, which are among the differen-

tial species for the sub-variation, are absent here, as

well as Vitex rehmannii, which is common elsewhere.

Several grasses which differentiate the Rhynchelytrum

vidosum-Schizachyrium jeffreysii community are

absent. They are: Diheteropogon amplectens, D.

filifolius, Schizachyrium jeffreysii, Elionurus argenteus,

Brachiaria serrata, B. nigropedata, Pogonarthria squar-

rosa and Urelytrum squarrosum. Within the community

Fimbristylis hispidula only occurs in three of the four

releves that typify the vegetation zones around old

settlements.

(c) Releve Nos 71, 73, 60, 111 and 208, representing

heterogeneous and disturbed quadrats, are grouped

together in Table 1.

1 . 1 . IB Transitional sub-variation

One of the three sub-variations transitional to the

variations on lower slopes fringes the northern

footslopes of Maroelakop, occuring on very slightly

sloping sites with eutrophic Hutton soils. The densely

wooded vegetation is typified by four releves, Nos.

171, 172, 169 and 170, of which no detailed structural

analysis was made.

An average of 74 species, including 19 woodies, was

found per plot. This is the highest average on the

Reserve.

The vegetation cover is usually between 1 and 5%

at each of the following height levels: 10-14 m;

8-10 m; 3-5 m; and a 0,5-3 m shrub layer. The

field layer is up to 1,75 m high with 30-65% cover.

Burkea africana, Terminalia sericea and Dichrostachys

cinerea subsp. glomerata, are the dominant woody

plants. Setaria perennis dominates the field layer and

generally has a high cover, indicating moist conditions

as in the Eragrostis pallens — Setaria perennis Variation

(1.1.2).

Common differential species in the field layer are

Felicia fascicularis, Stylosanthes fruticosa, Sporobolus

fimbriatus, Sphedamnocarpus pruriens, Ziziphus zeyhe-

rana. Acacia caff r a and Eragrostis acraea. The last

four species occur only in this sub-variation. Some of

the differential species are typical also of communities

on the heavy lowland soils. These species are Acacia

nilotica, A. karroo, A. tortilis, Ehretia rigida, Justicia

protracta and Achyropsis avicularis. Further relation-

ship with these communities is shown by the presence

of Dichrostachys cinerea subsp. glomerata, in the

tree and shrub layers; and Hibiscus engleri, Achyran-

thes aspera, Celosia linearis, Blepharis maderaspatensis

and Eragrostis lehmanniana in the field layer. Vernonia

oligocephala, Justicea anagalloides and Fadogia monti-

cola have relatively high presence as in the Eragrostis

pallens — Setaria perennis Variation (1.1.2).

This sub-variation is transitional to the Panicum

maximum — Acacia tortilis Communities of bottom-

land soils on the one hand, and the Eragrostis pallens —

Setaria perennis Variation of lower sandstone areas

on the other hand.

1 . 1 . 1C Transitional sub-variation

Another transitional sub-variation occurs on the

high sandstone areas where outcrops are common

and the soils are less deep and classified into the

Mispah Series. Apart from considerably more

Ochna pulchra shrubs and less Grewia flavescens shrubs

there is no marked difference in dominants, structure

and height with the typical sub-variation (1.1.1 A;

Fig. 7).

B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

149

Several species indicate a relationship with the two

remaining variations on moister, lower sandstone

areas (1.1.2 and 1.1.3). Such species are Trachy-

pogon spicatus, Schizachyrium sanguineum, Parinari

capensis, Pygmaeothamnus zeyheri and Arislida aequig-

lumis.

The sub-variation is transitional particularly to the

Eragrostis pollens — Trachypogon spicatus Variation

(1 . 1 .3) of the slightly sloping valley west of Maroela-

kop.

1 . 1 . 1 D Transitional sub-variation

On the middle slopes of the sandstone area where

soils gradually grade from the Hutton to the Clovelly

Form, a third transitional sub-variation occurs.

In dominants and structure the vegetation does not

differ clearly from the previous or from the typical

sub-variation (Table 2; Fig. 7). The same species

mentioned under the former sub-variation (1.1. 1C)

indicate relations with the remaining two variations

on lower slopes (1.1.2 and 1.1.3). The sub-variation

is particularly related to the Eragrostis pollens — ■

Setaria perennis Variation (1.1.2). Common features

are the constant high cover of Elionurus argenteus;

and the presence of Vernonia oligocephala and

Justicia anagalloides, which occur also in sub-variation

1 . 1 . 1 B fringing Maroelakop.

1.1.2 Eragrostis pallens — Setaria perennis Variation

This variation occurs in lower sandstone areas on

distrophic Clovelly soils (Mosdale Series) with

medium sand.

The vegetation differs from the Eragrostis pallens —

Dombeya rotundifolia Variation (1.1.1) of upper

slopes in the presence, height and cover of various

woody species (Table 2; Figs. 7 and 10).

Scattered individuals of Faurea saligna, up to 10 m

tall, occur. The dominants in the tree and shrub layers

are Burkea africana , Terminalia sericea and Ochna

pulchra. Eragrostis pallens and Setaria perennis are

dominant in the field layer.

The general height distribution of dominant woody

species is as follows:

Above 8 m: Burkea africana, followed by Termi-

nalia sericea.

5-8 m: Burkea africana, followed by Termi-

nalia sericea and Ochna pulchra.

3-3 ,5m: Terminalia sericea, followed by Burkea

africana.

2-3 m: Burkea africana, followed by Termi-

nalia sericea, Grewia flavescens and

Ochna pulchra.

1- 2 m: Ochna pulchra, followed by Burkea

africana and Terminalia sericea.

Below 1 m: Ochna pulchra, followed by Burkea

africana.

The average number of tree species per plot is seven,

which is low compared to the average of 14 in the

variation on upper slopes (1.1.1).

Total tree cover is considerably lower than in the

latter Eragrostis pallens — Dombeya rotundifolia Varia-

tion, usually less than 5%. Shrub cover is less than

10% and a field layer of up to 1,8 m high covers

30-75%.

Differential species are few and of low constancy.

They include Hypoxis rooperi and Tristachia rehmannii.

The high cover of Setaria perennis and Elionurus

argenteus is characteristic. Brachiaria serrata, Themeda

triandra and Chaetacanthus costatus have high

constancy.

1.1.3 Eragrostis pallens — Trachypogon spicatus

Variation

This variation occurs in the moderately sloping

valley west of Maroelakop on coarse textured meso-

trophic Clovelly soils (Sibakwe Series). The soils are

better drained than those of the former variation

because of their location at higher slope positions and

their coarse texture.

The vegetation resembles the previous variation in

presence, height and number of woody species.

However, Terminalia sericea has generally a higher

cover than Burkea africana at heights above 3,5 m.

Total tree cover is also greater in this variation and

can be up to 20%. Common woody species at different

heights are (Fig. 7 and Table 2):

Above 3 m: Terminalia sericea, followed by Burkea

africana.

2- 3 m: Burkea africana, followed by Termi-

nalia sericea.

1-2 m: Burkea africana and Ochna pulchra,

followed by Terminalia sericea.

Below 1 m: Ochna pulchra, followed by Burkea

africana and Terminalia sericea.

Fig. 10. — Physiognomy of the

Eragrostis pallens — Setaria

perennis Variation.

150

A PH YTOSOCIOLOGICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

A low average of seven woody species was recorded

per plot. Trachypogon spicatus and Eragrostis pallens

are dominant in a 1 ,75 m tall field layer with a total

cover of 30-50%.

A small group of species with low constancy

differentiates the variation. Conspicuous ones include

Terminalia phanerophlebia in the tree and shrub layer

and Cymbopogon plurinodis in the field layer. The high

cover of Trachvpogon spicatus is also characteristic.

1.2 Barleria bremekampii — Diplorhynchus Tree

Savanna (Table 3).

This community occurs on gentle to moderately

steep slopes of rocky sandstone hills and ridges, where

soils are litholitic and rocks cover 10-60% (Fig. 11).

The vegetation is broad-leaved savanna. Uneven,

broken areas carry a dense woody vegetation and on

more even terrain trees are less dense. Trees and

shrubs of varying age and height occur in all parts of

the savanna.

Common differential species include the tree

Diplorhynchus condylocarpon, the shrub Barleria

bremekampii and the forbs Tephrosia longipes,

Rhynchosia totta , Corchorus kirkii, Indigofera comosa.

Asparagus sounder siae and Euphorbia neopolycne-

moides. Another differential species, Xerophyta retiner-

vis, which occurs typically on very shallow soils and

amongst outcrops, is conspicuous because of its

unusual growth form of a protective cylinder of

moisture absorbing leaf sheaths round a thin woody

stem with adventitious roots (cf. Ayensu, 1973).

Three community variations correspond closely

with the main physiognomic differences and are

related to rockiness. Differences in aspect and degree

of slope are apparently of less importance.

1.2.1 Pseudolachnostylis — Diplorhynchus Variation

This savanna is typical of the Maroelakop area,

where it occurs on various aspects and degrees of slope,

with 30-60% rock cover.

The vegetation has a dense phase on uneven rocky

sites where trees are between 4 and 8 m high with

30-60% cover, and a more open phase with 4-6 m

tall trees covering 5-20%. Diplorhynchus condylo-

carpon is the only dominant tree. A 6, 5-2, 5 m shrub

layer covers up to 15% of quadrats with dense stands

of Canthium gilfillanii\ and a field layer of up to 1,50

m has 5-40% cover, depending on the amount of

rock. Schizachyrium jeffreysii is usually the dominant

grass and Setaria lindenbergiana is locally dominant.

Differential species include the tree Pseudolachno-

stylis maprouneifolia , the shrub Canthium gilfillanii

and the forb Polygala rehmannii. The habitat of the

relatively open phase of this community variation is

very similar to that of the Enneapogon scoparius —

Diplorhynchus Variation (1.2.2) on other rocky

sandstone areas.

1.2.2 Enneapogon scoparius — Diplorhynchus Varia-

tion

The moderately open savanna which covers most

of the relatively unbroken rocky slopes of Stemmers-

kop and the western sandstone ridges belongs to this

variation, which occurs on northerly and southerly

aspects with moderately steep slopes and 30-60%

rock cover.

The trees are mainly 4-6 m tall with 5-20% cover.

Shrub cover varies from 1-15% and grass and forb

cover is between 25 and 50%. Combretum molle and

Croton gratissimus are the dominant trees and

Enneapogon scoparius is the dominant grass.

Common differential woody species include Croton

gratissimus and the dwarf shrub Rhus magalismon-

tanum. Differential species in the field layer are

Enneapogon scoparius and the forbs Becium angusti-

folium and Vernonia suther/andii.

The characteristic presence and dominance of

Croton gratissimus and Enneapogon scoparius is

indicative of hot and dry conditions (cf. Theron, 1973;

Coetzee. 1974; and 1975).

1.2.3 Burkea africana — Diplorhynchus Variation

This variation occurs on gentle slopes with 10-20%

outcrop. Trees are between 4 and 6 m high and

usually have up to 5% cover, which is much lower

than in the previous variations on rockier sites.

Shrubs cover up to 5 % of quadrats, while grasses and

Fig. 1 1 . — Barleria bremekampii —

Diplorhynchus Tree Savanna

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B J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

151

forbs have 50-60% cover — the highest in the com-

munity. Burkea africana and Diplorhynchus condylo-

carpon are the dominant trees and Loudetia flcivida and

Schizachyrium jeffreysii the dominant grasses.

Species which differentiate the community variation

on their characteristically high cover are Burkea

africana, Brachiaria serrata, Loudetia flavida and

Merremia tridentata.

1.3 Eragrostis racemosa — Schizachyrium jeffreysii

Tree Savanna and Grassland (Table 3).

This community includes all the felsite elevations

and the footslopes of the southern sandstone areas

and covers approximately a third of the Reserve. The

topography is gently sloping to virtually level and

during winter nights frost intensity presumably varies

from mild on upper slopes to more severe on lower

slopes. Because of their felsite origin and lower slope

position the sandy soils are richer in minerals than

soils of related communities on sandstone (1.1 and

1.2).

Broad-leaved savanna occurs on the uplands, where

shallow, well-drained soils are underlain by weathered

felsite. Grasslands occur on frostier lower slopes with

shallow to deep poorly drained soils, underlain by a

hard flintite bank. Most of the deeper grassland soils

have mottled perched gley horizons and soft plinthic

B-horizons, a result of fluctuating water tables and

lateral movement of water. The termitaria thickets

that occur throughout these savannas and grasslands

belong to a different community (2 . 2).

Differential species include 14 grasses and eight

forbs. Table 3 shows that the most common differen-

tial grasses, Elionurus argenteus, Trachypogon spicatus,

Setaria perennis and Brachiaria serrata, are not

exclusive to this community. Exclusive differential

grasses are Eragrostis racemosa, E. gummiflua and

Digitaria monodactyla. Constant differential forbs are

Agathisanthemum bojeri, Nidorella hottentottica and

Stylosanthes fruticosa.

Grunow (1965) described related savanna and

grassland communities from Soutpan, near Pretoria.

The distinctive habitat feature of the grassland there is

soil drainage impeded by an ironpan near the soil

surface. On the poorly drained felsite areas at Nylsvley

frost appears to be an additional habitat factor

favouring grasslands.

1.3.1 Eragrostis racemosa — Combretum apiculatum

Tree Savanna.

Most of this savanna community on felsite occurs

on the upper slopes of a broad ridge in the north-

west and on a small isolated eastern elevation.

Both stands occur uninterruptedly and coincide with

soils of the Mispah Series (Mispah Form), which are

shallow, well-drained and underlain by weathering

felsite. Parts of the community occur on litholitic

soil and on a local patch of soil belonging to the

Glenrosa Series (Gemvale Form), which has a coarse

to medium textured sandy orthic A-horizon and a

cutanic B-horizon with 15-30% clay.

Tree density in the felsite savanna varies consider-

ably. Open stands, with approximately 20% tree

cover, are the most extensive. Trees become sparse

with less than 5% cover towards the grasslands and

dense tree stands with up to 60% cover occur in belts

throughout the savanna region. These dense stands

are presumably situated above interbedded rocks with

favourable moisture conditions. This was suggested by

Theron (1973) as a tentative explanation for the

occurrence of similar dense belts in the felsite savannas

at Loskopdam. Grass and forb cover in the savanna

is usually between 40 and 70%.

Combretum apiculatum and Vitex rehmannii are

common trees in all three savanna phases and are also

the most conspicuous differential species of the

community. Other differential species include the

low shrub Lippia javanica and the forbs Ruellia

patula and Crabbea angustifolia.

The felsite savanna has a group of species in

common with communities of sandstone uplands

(1.1 and 1 .2). The characteristic species combination,

particularly that of the typical Rhus leptodictya—

Combretum apiculatum Variation (1.3. 1.1), shows

affinities also with the Euclea undulata — Acacia

tort i/is Tree Savanna of the bottomlands (2.2).

Theron (1973) found a highly significant positive

regression of Combretum apiculatum, which is one of

the differentiating species of the felsite savanna, with

iron content of the soil. The closely related Andropogon

amplectens — Combretum apiculatum — Bulbostylis col-

lina Community described by Grunow (1965) occurs

on very shallow, sandy, acid, sedentary soils, underlain

by iron pan. A typical variation and a transitional

variation to grassland were found.

1.3. 1.1 Rhus leptodictya — Combretum apiculatum

Variation (Fig. 12)

The extensive typical variation includes the mod-

erately open to dense savannas, with 20-60% tree

cover. Most trees fall in the 2-6 m height range and

several species have individuals of various ages over

the entire height range. The tallest trees, which are

sparsely scattered, are usually up to 8 m and occasion-

ally up to 10 m high.

The typical variation has more than 20 woody

species of which 17 occur in at least 25% of the

releves. The dominant woody species are Combretum

apiculatum, Vitex rehmannii, Combretum mode, Pelto-

phorum africanum, Terminalia sericea and Burkea

africana. The dominant grasses are usually one or

more of the following: Setaria perennis, Laudetia

flavida, Themeda triandra, Trachypogon spicatus,

Rhynchelytrum villosum, Schizachyrium sanguineum

and Elionurus argenteus.

Nearly all differential species are woody and of these

the following have high constancies: Rhus leptodictya,

Pe/tophorum africanum, Euclea undulata and Terminalia

sericea.

1 . 3 . 1 . 2 Cymbopogon plurinodis — Combretum api-

culatum Variation

This variation, represented by 5 releves, is tran-

sitional to Eragrostis racemosa — Digitaria monodactyla

Grassland (1.3.2) on frosty lower slopes with poorly

drained soils. Woody plants have less than 5% cover

and include sparsely scattered tall trees, interspersed

with small trees and shrubs that are predominantly

between 1 ,5 and 4 m in height.

The transitional variation has much fewer constant

woody species than the previous typical variation.

Woody species occurring in more than one releve

are: Vitex rehmannii, with 100% presence; Combretum

apiculatum (80% presence); Acacia caffra (60%);

Combretum zeyheri (40%); and Parinari capense, a

low shrub in the field layer, with 40% presence. The

dominant shrubs are usually either Vitex rehmannii or

Acacia caffra. Dominant grasses include Elionurus

argenteus, Digitaria cf. eriantha, Cymbopogon pluri-

nodis, Trachypogon spicatus, Setaria perennis, Eragros-

tis curvula and Schizachyrium sanguineum.

152

A PHYTOSOCIOLOGICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

1.3.2 Eragrostis racemosa — Digitaria monodactyla

Grassland

The grassland community of relatively frosty and

poorly drained areas occurs on slopes at the bottom

end of a broad valley in the northernmost part of the

Reserve and as a narrow fringe on the lowest slopes of

the sandstone and felsite areas (Fig. 13).

Cover in the grassland varies from 50-80%. A

short, xeric grassland variation occurs on very shallow

dry soils of convex slopes and a tall mesic grassland

variation occurs on deep soils of concave and flat

areas where soil and moisture accumulate.

Although the community has few positive differen-

tial species, it is strongly negatively defined. The most

constant differential species are the geophyte Lede-

bonria revoluta and the grass Aristida curvata.

1.3.2. 1 Eragrostis nindensis — Digitaria monodactyla

Variation

The xeric short grassland variation occupies most of

the predominantly convex slopes in the northern

part of the Reserve. The soils are shallow, gravelly

and stony, usually dry but underlain by a hard

impenetrable flintite bank causing impeded soil

drainage during wet periods. Elionarus argenteus

is the only constant dominant. Occasional co-domi-

nants include Setaria perennis, Loudetia flavida,

Rhynchelytrum vil/osum, Digitaria cf. eriantha, Trachy-

pogon spicatus, Brachiaria serrata and Aristida diffusa.

In the xeric grasslands, besides numerous termitaria

bush clumps, one also finds isolated tall trees or small

groups of trees. These are mainly Combretum apicu-

l at um. Acacia caffra, Peltophorum africanum and

Burkea africana. Near alluvial areas intrusive shrubs of

Acacia tor til is, A. nilotica and A. karroo may occur

in the grasslands.

The community is differentiated by Hyperthelia

dissoluta and by Eragrostis nindensis, a desiccation

tolerant grass, typical of xeric grassland (cf. Gaff &

Ellis, 1974; and Coetzee, 1975).

Fig. 12. — Rhus leptodictya —

Combretum apiculatum Va-

riation on felsite uplands.

Fig. 13. — Eragrostis racemosa —

Digitaria monodactyla Grass-

land with scattered termi-

taria thickets.

B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

153

1.3. 2. 2 Tristachya rehmannii — Digitaria mono-

dactyla Variation

The mesic tall grassland variation occurs on deep,

non-gravelly and moist soils found mainly on flat

to concave, very gentle slopes in areas of soil and

water accumulation. Included are parts of the felsite

valley in the north and the grassland fringes on te

lowest felsite and sandstone slopes. The deep grass-

land soils of the felsite region belong mainly to the

Wasbank and Longlands Forms, which have orthic

A-horizons; perched gley horizons, mottled with

accumulated iron and magnesium oxides; and plinthic

B-horizons — a type of profile resulting from prolonged

poor drainage. The deep soils of the sandstone grass-

land belong mainly to the Fernwood Series (Fernwood

Form) and Soetmelk Series (Avalon Form). The

Fernwood Series is an orthic A-horizon on medium

textured regie sand and the Soetmelk Series comprises

an orthic A-horizon, yellow apedal B-horizon and a

soft plinthic B-horizon with 15-35% clay.

Tristachya rehmannii, a tall grass with very high

fibre content, and Setaria perennis, a typical meso-

phytic grass (see also 1.1.2), are characteristically

dominant. Occasional co-dominant and sub-domi-

nants include Trachypogon spicatus, Elionurus argen-

teus, Digitaria cf. eriantha and Eragrostis gummiflua.

Differential species are Tristachya rehmannii (a

dominant) and the mesophytic forb Helichrysum

nudifolium.

2. Communities of flat bottomlands and of termitaria

{excluding the self-mulching and vertic soils)

(Table 4)

(Panicum maximum— Acacia tortilis Tree

Savannas and Termitaria Thickets)

Part of the flat bottomlands consists of recent

alluvium with a variety of calcareous clay soils. The

vegetation is microphyllous thorn savanna. Included

also are termitaria thickets which occur throughout

the Reserve except on the sandstone elevations. Tree

density varies from canopies interlocking on the

termitaria to sparsely wooded grassland on very

brackish clay soils. Trees vary in height up to 10 m,

the tallest trees occurring on termitaria. The field

layer varies in height and cover depending on soil

brackishness and the amount of grazing. Common

differential species are Acacia tortilis in the tree and

shrub layer; and Panicum maximum. Aloe spp. and

Chloris virgata in the field layer.

Floristically related communities were described

by Grunow (1965) from the farm Soutpan where

Acacia tortilis is very common in communities on

deep, well-drained, alkaline soils.

The savanna has two communities associated with

differences in soil texture and soil brackishness.

2. 1 Sporobolus ioclados — Acacia tortilis Tree Savanna

This community occurs on very heavy, compact,

hard and brackish soils, belonging to the Shephardvale

Series (Valsrivier Form; Fig. 14). The soils have a

strongly structured B-horizon with more than 55%

clay. All plots were heavily grazed and poor in

species, the average number of species being 19 per

plot. The vegetation usually consists of 2-6 m high

trees, covering up to 10%; a 0,5-2 m shrub layer with

less than 1 % cover; and a field layer up to 0,5 m with

15-50% cover.

The community is strongly differentiated by

Sporobolus ioclados and Ocimum canum. The tree

Boscia foetida var. rehmannii is also characteristic,

but occurs sporadically.

2.2 Euclea undulata — Acacia tortilis Tree Savanna

and Termitaria Thickets

This community includes the vegetation on less

brackish, less heavy and less compact soils and on

termitaria of the bottomlands. The vegetation varies

from dense, impenetrable thicket on the termitaria to

open savannas on alluvium. Carissa bispinosa and

Euclea undulata are dominant and differential woody

species. Two variations are recognized.

Fig. 14. — Sporobolus ioclados —

Acacia tortilis Tree Savanna

on heavy brackish soils.

Acacia tortilis in left fore-

ground.

154

A PHYTOSOCIOLOGICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

TABLE 4. Communities of termitaria and alluvial flats.

D = differential for one community type; d = differential for

two or more community types.

Species of double and single occurrence

Abutilon austro-africanum (205sr; 198:r), Acacia robusta (132:1; 205:+), Asparagus africanus

(160: + ; 195:r), Boscia foetida (I57sr; 128:r), Bothriochloa insculpta (132:+; 205'-+), Conyza

bonariensis (196:+; 195:+), Crotalaria sphaerocarpa (l60:+; r?4:r), Eragrostis obtusa (206:+;

195*+), Eragrostis curvula (195:+; 205:r), Eragrostis rigidior (206:1; 198:1), Rhus lancea

(l30:r; 131:r), Scolopia zeyheri (130:2; 131: r), Securinega virosa (l33;r; 205:+), Tarchonanthus

camphoratus (129 : + ; 198:2), Vernonia poskeana (l57:r; 194:r).

Acacia hebeclada (128:2), Achyranthes sicula (128:+), Acrotome angustifolia (l57:r),

Artemieiopsis villosa (158:1), Asparagus virgata (l98:r), Bidens bipinnata (l96:r), Bidens

pilosa (194:+), Bonatea antennifera (l31:r), Crabbea hirsuta (205:+), Denekia capensis ( 159 J +) *

Dicliptera micranthe6 (l96:r), Dovyalis zeyheri (130:2), Echinochloa holubii (157:+), Epaltes

gariepina (159*1) , Eragrostis ciliarensis (l32:r), Ereigrostis trichophora (157*2), Erigeron

floribundus (133* r), Justicia natammensis (l60:r), Lepidium africanum (l96:r), Leucas

neufliziana (l74:r), Monsonia angustata (l74:r), Monsonia biflora (l60:r), Nidorella resedifolia

(l98:r), Otiophera sp. (l57*r), Panicum deustum (198:+), Panicum laevifolium (198:+), Portulaca

pilosa (174:+), Rhus gueinzii (206:+), Ruschia sp. (129*+), Scabiosa columbaria (198:+),

Schkuhra pinnata (l31:r), Schmidtia pappophoroides (174:1), Scirpus sp. (194:+), Sporobolus

iocladoo (194:+). Sporobolus panicoides (l28:r), Striga gesnerloides (157*+), Vernonia

faacicularia (198:r).

B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER

155

2.2.1 Pappea capensis — Acacia tortilis Variation

This variation comprises the vegetation on termi-

taria of the felsite elevations. There are many woody

species. Plant growth is dense and sometimes virtually

impenetrable because trees, shrubs and climbers

interlock (Fig. 15).

A 2-8 m tree and tall shrub layer has 30-70%

cover. Lower strata are: a 0,5-2 m shrub layer with

5-25% cover; and a field layer up to 0,5 m with 10-

40% cover. Euclea undulata and Ziziphus mucronata

dominate the tree and tall shrub layer and Carissa

bispinosa dominates the low shrub layer. The most

common differential woody species is Pappea capensis.

In the field layer Kalanchoe lanceolata, Plectranthus

cylindraceus and P. neochilus are typical differentials.

2.2.2 Acacia nilotica — Acacia tortilis Variation

(Fig. 16)

This variation occurs typically on calcareous

alluvium. The relatively sandy soils on the river

banks belong to the Limpopo Series (Oakleaf Form)

and the heavier soils belong to the Shephardvale

Series (Valsrivier Form). Part of the variation is

found on dark calcareous clay soils with vertic

characters and crusting surface belonging to the

Gelykvlakte Series (Arcadia Form).

Acacia tortilis is the dominant tree. Carissa bispinosa

dominates the shrub layer, and Panicum maximum and

Eragrostis lehmanniana dominate the field layer.

The vegetation consists of a 6-10 m tree layer with

up to 10% cover; a 2-5 m tree layer with 1-5 % cover;

a 0,5-2 m shrub layer with 1-5% cover; and a 1 ,3 m

high field layer with 20-75% cover, depending on the

amount of grazing. Common differential species are

Acacia nilotica, A. karroo and Blepharis transvaalensis.

Releve Nos. 206, 156 and 193 are from bottomland

termitaria and are transitional between the bottom-

land alluvium variation and the upland termitarium

variation. Several differential species of the upland

termitaria are present, including Cassine transvaalensis,

Jasminum breviflorum and Maytenus heterophylla,

but others such as Pavetta assimilis, Plectranthus

neochilus and P. cylindraceus are absent. This illu-

strates a floristic gradient from elevated felsite

Fig. 15. — Termitarium thicket

belonging to the Pappea

capensis — Acacia tortilis Va-

riation.

Fig. 16. — Acacia nilotica — Acacia

tortilis Variation on alluvium.

156

A PHYTOSOCIOLOG ICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

termitaria thicket (2.2.1), via bottomland termitaria

thicket, to alluvium savanna (2.2.2). This gradient is

associated with special termitarium conditions (Van

der Schijff, 1957) and water table.

A separate alluvial community, of small extent,

occurs south of the main road to Holme Park. The area

is flat and low-lying with yellowish brown silty clay

loam soil. The vegetation consists of a 2-3 m tall

woody stratum with 2% cover, dominated by Rhus

pyroides and Diospyros lycioides and a field layer

with 75% cover, mainly of Elionurus argenteus and

Eragrostis spp. Sparsely scattered tall Acacia mellifera

subsp. detinens and Ziziphus mucronata trees and

occasional small Acacia tortilis shrubs also occur.

The woody species indicate relationships with other

alluvial savannas but the field layer is distinctly

different with elements of marshy grasslands and

communities of self-mulching vertic soils. The

following is an example of this Rhus pyroides —

dominated savanna:

Re lev e No. 216 (10x20 m)

Trees and shrubs:

Rhus pyroides 1

Acacia tortilis +

Acacia mellifera subsp. detinens r

Diospyros lycioides r

Field layer:

Denekia capensis +

Epaltes gariepina +

Bergia decumbens +

Elionurus argenteus 2 b

Eragrostis atrovirens +

Panicum color atum +

Eragrostis plana +

Cyperus sphaerospermus +

Setaria sphacelata +

Conyza bonariensis +

Verbena bonariensis +

Commelina africana +

Fimbristylis dichotoma +

Amphidoxa filaginea +

Nidorella resedifolia +

3. Communities of self-mulching vertic soils

(Aristida bipartita — Setaria woodii Tree Savanna

and Grassland', Fig. 17)

The self-mulching vertic soil belt of the Arcadia

Series (Arcadia Form) occurs in the south-eastern

bottomland bordering the elevated sandstone area.

Habitat features include a fluctuating water table,

prolonged periods of inundation during heavy

rainfall, swelling and contracting of the soil during

wet and dry periods with considerable cracking when

dry, a loose soil surface, a high calcium carbonate

content in the soil and gilgai micro-relief.

Fig. 17. — Vertic black clay soil of the Aristida bipartita — Setaria

woodii Tree Savanna and Grassland.

TABLE 5. Communities of self-mulching, vertic clay.

D = Differential for one community type; d=differential for two or more community types.

Species of single occurrence:

Acanthosicyos naudiniana (168: + ), Asparagus suaveolens (I97:r), Celosia linearis ( 1 64 :r), Epaltes gariepina (161:+), Eragrostis

heteromera (168:+), Mariscus sp. (162: + ), Maytenus heterophylla ( 1 97 :r), Nidorella resedifolia (199:r), Panicum natalense (168: + ),

Psoralea holubii (162: + ), Rhus pyroides (200:r), Securidaca longipedunculata (168: + ), Tephrosia lupinifolia (168:+), Wahlenbergia

caledonica ( 1 68 :r).

157

B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WE1SSER

The vegetation is mainly grassland but an open

stand of thorn savanna occurs in the slightly higher

north-western part where the water table is generally

lower than in the grassland. Low termitaria mounds,

several metres in diameter, also occur here. These

better drained mounds have thickets belonging to the

Acacia nilotica — Acacia tortilis Variation (2.2.2).

The field layer of the savanna phase corresponds to

the grassland phase and this stratum has very few

species in common with other communities in the

Reserve. Grass and forb cover is between 80 and 90%

and dominants include Setaria woodii, Dichanthium

papillosum , Scirpus dregeanus and Aristida bipartita.

The grass stratum is a mosaic of varying dominants

coinciding with crests and troughs in the gilgai micro-

relief. Aristida bipartita is typically dominant on the

relatively xeric crests. Tree cover in the savanna is

between 1 and 5 %. The trees, mostly Acacia karroo ,

vary in height from small juveniles to 9 m tall. Zizi-

phus mucronata is the sub-dominant tree.

Trees have been cleared from part of the savanna

area. Seedlings are common in parts of the grassland,

but extend further into the grassland than from where

the trees were cleared. We doubt that the occurrence

of the seedlings indicates the potential range of the

savanna, because the seedlings in the lower lying parts

are probably periodically drowned. Interference with

the water table level may, however, result in extensive

savanna and thicket encroachment at the expense

of pure grassland.

The two components in the field layer mosaic may

be separately characterized as has, e.g., been done by

Van Zinderen-Bakker & Werger (1974) in the high

altitude bogs of Lesotho and by Verster, De Villiers &

Scheepers (1973), in closely related gilgai vegetation

on soils of the Arcadia Series, near Rustenburg.

Verster et al. (1973) found that numerous species

favoured either crests or troughs and that crests are

floristically richer than troughs. In the present study

each releve covers both components. Until recently,

part of the self-mulching vertic soils have been

ploughed to produce fodder crops. The present

secondary vegetation differs clearly from the sur-

rounding grassland community The following is an

example:

Rcdeve No. 215 (lOx 10 m)

Total cover of grasses and forbs: 60-65%.

Dinebra retroflexa 4

Aristida bipartita 4-

Setaria woodii r

Brachiaria eruciformis -\-

Psoralea hohibii r

Sutera sp -f-

Senecio apiifolius +

Phyllanthus cf. maderaspatensis

4. Communities of abandoned settlements (Table 6)

On the elevated sandstone area, the disturbance

effect on the vegetation around previous scattered

native settlements can still be seen. The eutrophic

Hutton soils here have a high phosphate content.

The vegetation is differentiated by Eragrostis lehman-

niana, which is the only dominant grass, sometimes

with up to 80% cover and by Solanum delagoense

and Crotolaria pisocarpa. Occasional tall Sclerocarya

caffra trees around the settlements are characteristic.

The former differential activities of the tribes are

still reflected in the present vegetation. Thorn trees,

mainly Acacia tortilis, occur on housing and living

sites where the soils are hard and compact (Fig. 18).

Sparsely wooded grassland dominated by Eragrostis

lehmanniana occurs on the looser soils where agricul-

ture must have been practised.

The thorn tree vegetation consists of a 2-6 m tree

layer with up to 10% cover; a 0,5-2 m shrub layer

with up to 5% cover; and a field layer up to 0,8 m

high with 25—95 % cover. Schkuhria pinnata, Leucas

neuflizeana and Eragrostis rigidior are typical species

in the field layer.

The grasslands have 0,5-3 m high scattered trees

and shrubs, mainly Burkea africana and Terminalia

sericea; and a field layer of up to 0,5 m with 40-80%

cover. Cassia absus, Stipagrostis uniplumis, Eragrostis

superba, Corchorus asplenifolia and Aristida stipitata

are typical field layer species.

Fig'. 18. — Acacia nilotica and

Acacia tortilis trees on aban-

doned settlement.

158

A PHYTOSOCIOLOGTCAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

TABLE 6. Secondary communities.

D = Differential for one community type; d = differential for

two or more community types.

Differential species

D Eragrostis lehmanniana

D Solanum delagoense

D Crotolaria pisocarpa

Differential species of Variation 4.1

D Cassia absus

D Stipagrostis uniplumis

D Eragrostis superba ....................

D Corchorus aspleni folia

D Aristida stipitata

Differential species of Variation 4.2

D Schkuhria pinnata

D Leucas neufliziana

D Eragrostis rigidior

D Cynodon dactylon

D Portulaca quadrifida ..................

D Corchorus tridens

Species common to primary and secondary

community types

d(l.l) Vernonia poskeana

d ( 1 ) Evolvulus alsinoides

d(l) Heteropogon contortus

d ( 1 ) Walteria indica .......................

d(l) Tephrosia burchellii ..................

d(l) Digitaria eriantha

d(l.l.lA;2) Panicum maximum

d(l.l) Borreria scabra

d(l.l) Cleome rubella

d(l.l) Limeum viscosum

d(l.l) Ehynchosia longiflora

d(l) Rhynchelytrum villosum

d ( 1 ) Burkea africana

d(l) Terminalia sericea

d(l.l) Striga asiatica

d(l) Indigofera daleoides

d(l) Kohautia lasiocarpa

d(2;l.l.lB) Acacia tortilis .......................

d(see 2.2.2)Acacia nilotica .......................

d(l.l) Sida cordifolia

General and infrequent species

Aristida congesta

Dichrostachys cinerea subsp. glomerata

Achy rant he s aspera ....................

Monsonia angustata

Sesamum alatum

12 3 1

r + r r

1 + +

3 3

+

1

3 2 5 2 2 3

2 12 + 12

+ 1 + + 1 2

r + r + + r

3 2 + + +

1 r +

+ 21

+ + r

r r r + r r

112

+ 1 +

r r r r r r

+ r + r +

Species of double and single occurrence

Acacia karroo (31:r; 173;r), Aristida adscensionis (32:1; 173:+),

Chloris virgata (32:+; 174:+), Crotolaria sphaerocarpa (l74:r; l84:r),

Enneapogon scoparius (32:+; 33:r), Indigofera macra (22:r; 31;r),

Pollichia campestris (l73;r; l84:r), Schmidtia pappophoroides (l73:+i

174:1), Senecio apiifolius (32:r; 33:+), Sida alba ( 173 : + ; l84:r).

Abutilon austro-africanum (l84:r), Acacia caffra (34:r),

Alternanthera pungens (32:2), Amaranthus thunbergii (32:r), Aristida

canescens (32:r), Asparagus plumosus (32:r), Asparagus suaveolens

(I73;r), Bidens pilosa (l84:+), Blepharis integrifolia (l84:+), Cenchrus

ciliaris (32:+), Chaetacanthus costatus (l84:r), Chenopodium carinatum

(32:+), Cleome monophylla (31:r), Convolvulus sagittatus (22:+),

Cymbopogon excavatus (22:2), Dichapetalum cymosum (I73;r), Dicoma

macrocephala (l84:r), Euphorbia inaequilatera (22:r), Euphorbia tirucalli

(173:+), Felicia fascicularis (l84:l), Felicia mossamedensis (33:+),

Gazania krebsiana (22:r), Gomphrena celosioides (173:+), Helichrysum

argyrosphaerum (32:+), Hibiscus cannabinus (174:+), Hibiscus engleri

(173:+), Hibiscus trionum (32:r), Indigofera rhytidocarpa (30:r),

Justicia flava (l84:+), Melhania transvaalensis (33;r), Orthanthera

jasminiflora (33:r), Panicum natalense (173:+), Polygala hottentotta

(22:r), Portulaca pilosa (174:+), Ehynchosia confusa (22:r),

Rhynchelytrum repens (22:+), Sclerocarya caffra (l84:l), Talinum caffrum

(30:r), Urochloa brachyura (32:+), Zinnia multiflora (33:r), Zornia

capensis (173:+)-

B. J. COETZEE, F. VAN DER MEULEN, S. ZWANZIGER, P. GONSALVES AND P. J. WEISSER 159

Many species occurring in this secondary vegetation

indicate relationships with the less disturbed sur-

rounding vegetation.

PHYSIOGNOMIC SYNOPSIS AND GENERAL REMARKS

The major physiognomic, dominance and physio-

graphic types in the Reserve correspond well with

floristically defined communities and can readily be

delineated in the field and on aerial photographs in a

precursory survey (Fig. 19).

1. Broad-leaved savannas are restricted to elevated

areas. Together with upland grasslands they form

one broad, floristically defined community

associated with sandy soils. The major floristic

sub-divisions of the community correspond to:

1 . 1 broad-leaved savanna on well drained, non

litholitic sandstone soils ( Burkea-Terminalia -

Ochna-Grewia dominated savanna);

1 .2 broad-leaved savanna on litholitic sandstone soils

( Diplorhynchus , or Burkea-Diplorhynchus, or

Combretum mol/e-Croton gratissimus dominated

savannas);

1.3 broad-leaved savannas on felsite [ Combretum

apiculatum-Vitex dominated savanna (1.3.1)];

and grassland on sandy soil [grassland dominated

by one or more of the following: Elionurus

argenteus, Setaria perennis, Loudetia ftavida,

Rhynchelytrum villosum, Trachypogon spicatus,

Brachiaria serrata, Aristida diffusa and Tristachya

rehmannii (1.3.2)].

2. Microphyllous thorn savannas on bottomland

alluvium and thickets on termitaria form another

broad floristically defined community, with sub-

divisions corresponding to:

2 . 1 sparsely wooded grassland on compact, very

brackish soils, with a field layer dominated by

Sporobolus ioclados\

2.2 thickets on well drained upland termitaria

(2.2.1); thickets on poorly drained bottomland

termitaria (2.2.2) and thorn savanna on rela-

tively loose alluvial soils, with a field layer

dominated by Panicum maximum and Eragrostis

lehmanniana.

3. Grassland and microphyllous thorn savanna of

self-mulching vertic soils form a broad community

which is divided into thorn savanna of the better

drained parts, and grassland dominated by

Setaria woodii, Dichanthium papillosum , Scirpus

dregeanus and Aristida bipart it a on the poorer

drained parts.

4. Secondary grasslands and thorn savannas occur

on abandoned lands and settlements.

The major physiognomic vegetation characters that

change in relation to floristically defined communities

are presence, cover and leaf or leaflet size of trees and

shrubs. Leaf or leaflet size is strongly coupled with the

presence or absence of thorns. The full floristic

analysis showed the total floristic composition,

characteristic species, floristic interrelationships, hier-

archical level, variability and finer sub-divisions of the

various community types and the phytosociological,

ecological and geographical distribution of each

species recorded in the survey.

The vegetation differences correspond well with

differences in soil forms, series and phases. Important

soil differences that are related to physiognomic and

floristic vegetation differences are texture, depth,

drainage and nutrient status as determined by geolo-

gical origin, topographic position and disturbance.

Fig. 19. — Schematic representation of main vegetation — habitat relationships at Nylsvley.

160

A PHYTOSOCIOLOG ICAL CLASSIFICATION OF THE NYLSVLEY NATURE RESERVE

Frequency and severity of frost, which are correlated

with topographic features and thus with soil diffe-

rences, may also be an important determining factor.

ACKNOWLEDGEMENTS

Mrs I. C. van der Meulen greatly assisted us during

the earlier phases of the project. Dr J. W. Morris helped

with the computer printing of tables, which were then

typed by Mrs J. M. Mulvenna. Mrs L. Williams gave

technical assistance and our colleagues in the herba-

rium named numerous plant specimens. Prof. H. J.

vonM. Harmseand Mr P. J. Botha provided informa-

tion on the soils. These contributions, as well as

valuable comments particularly by Dr D. Edwards and

also by Messrs H. C. Taylor, E. A. Galpin, K.

Hoffman, M. C. Rutherford and J. C. Scheepers, are

gratefully acknowledged.

UITTREKSEL

Die plant egroei van die Nylsv/ey-natuurreservaat in

die Transvaalse Gemengde Bosve/d is hierargies geklas-

sifiseer met behulp van die Braun- Blanquet Metode.

Die plantegroei is seisoenale grasveld en bladwisselende

savanne met vier floristies-gedefinieerde hoofgroepe

plantegemeenskappe :

(1) grasveld en breeblaar savannes op suur sanderige

gronde van hoogliggende sandsteen en felsietgebiede ;

(2) mikrofiele doringboomsavannes op kalkryke

kleierige alluviate gronde van 'n laagte en digte

termitaria-boskolle ;

(3) grasveld en doringboomsavanne op kalkryke

selfkrummelende vertiese grond ; en

(4) sekondere plantgemeenskappe op vroeere Bantoe-

nedersettings en onlangs geploegde grond.

Sewe primere gemeenskappe met 12 variasies en 4

subvariasies, en drie sekondere gemeenskappe word op

basis van 216 re/eves beskryf 'n Ekosisteemstudiegebied

is in meer detail as die res van die Reservaat bemonster

en beskryf.

REFERENCES

Acocks, J. P. H., 1953. Veld Types of South Africa. Mem. Bol.

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

Ayensu, E. S., 1973. Biological and morphological aspects of the

Velloziaceae. Biotropica 5: 135-149.

Coetzee, B. J., 1974. A phytosociological classification of the

vegetation of the Jack Scott Nature Reserve. Boihatia 1 1 :

329-347.

Coetzee, B. J., 1975. A phytosociological classification of the

Rustenburg Nature Reserve. Bothalia 11,4: 561-580

Du Plessis, J. C., 1972. ' n Floristies-ekologiese studie van die

plaas Doornkop in die distrik Middelburg , Transvaal. M.Sc.

thesis, Univ. Pretoria.

Edwards, D., 1974. Conservation areas in relation to Veld

Types. Koedoe: In preparation.

Galpin, E. E., 1926. Botanical survey of the Springbok Flats.

Mem. Bot. Surv. S. Afr. 12: 1-100.

Gaff, D. F. & Ellis, R. P., 1974. South African grasses with

foliage that revives after dehydration. Bothalia 1 1 : 305-308.

Grunow, J. O., 1965. Objective classification of plant communi-

ties: a synecological study in the Sour-mixed Bushveld of

Transvaal. D.Sc. (Agric.) thesis, Univ. Pretoria.

Grunow, J. O., 1974. Savanna ecosystem research project :

General information on Nylsvley farm. Unpubl.

Rutfierford, M. C., 1972. Notes on the flora and vegetation of

the Omuverume Plateau — Mountain, Waterberg, South

West Africa. Dinteria 8: 3—55.

Schulze, B. R., 1947. The climates of South Africa according to

the classifactions of Koppen and Thornthwaite. S. Afr

Geogr. J. 29: 32-42.

Schulze, B. R., 1965. Climate of South Africa. 8. General survey.

Pretoria: Govt. Printer & Weather Bureau.

Theron, G. K., 1973. ’// Ekologiese studie van die plantegroei van

die Loskopdamnatuurreservaat. D.Sc. thesis, Univ. Pretoria.

Trigonometrical Survey Office, 1967. South Africa 1:50000

sheet: 2428 DA Naboomspruit. Pretoria: Trig. Surv. Office.

Van Der Schijff, H. P., 1957. 'n Ekologiese studie van die

flora van die Nasionale Krugerwildtuin. D.Sc. thesis, Univ.

Potchefstroom.

Van Zinderen Bakker, E. M. & Werger, M. J. A., 1974.

Environment, vegetation and phytogeography of the high-

altitude bogs of Lesotho. Vegetatio 29 : 37-49.

Verster, E., Df. Villiers, J. M. & Scheepers, J. C., 1973.

Gilgai in the Rustenburg area. Agrochemophysica 4: 57-62.

Von M. Harmse, H. J. & Botha, P. J. Report on the soils of

Nylsvley Reserve. Pers. Comm.

Walther, H. & Lieth, H., 1960. Klimadiagramm — Weltatlas.

Jena: Fisher.

Weather Bureau, 1954. Climate of South Afrjca. 1. Climate

Statistics. Pretoria: Govt. Printer & Weather Bureau.

Weather Bureau, 1957. Climate of South Africa. 4. Rainfall

maps. Pretoria: Govt. Printer & Weather Bureau.

Weather Bureau, 1960. Climate of South Africa. 6. Surface

winds. Pretoria: Govt. Printer & Weather Bureau.

Weather Bureau, 1965. Climate of South Africa. 9. Average

monthly rainfall up to the end of 1960. Pretoria: Govt.

Printer & Weather Bureau.

Werger, M. J. A., 1974a. On concepts and techniques applied

in the Ztirich-Montpellier method of vegetation survey.

Bothalia 1 1 : 309-323.

Werger, M. J. A., 1974b. Applicability of Ziirich-Montpellier

Methods in African and sub-tropical rangelands. In R.

Tiixen (gen. ed.), Handbook of vegetation science: W.

Krause (ed.), 13 Application to grassland husbandry. The

Hague: Junk.

Westhoff, V. & Van Der Maarel, E., 1973. The Braun-

Blanquet Approach. Tiixen, R. (gen. ed.). Handbook of

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classification of vegetation. The Hague: Junk.

Bothalia 12, 1 : 161—165 (1976)

Thuranthos: notes on generic status, morphology, phenology

and pollination biology

C. H. STIRTON*

ABSTRACT

The generic status of Thuranthos C. H. Wr. is discussed. A key separating the two species is provided. The

genus is reported to occur in Natal for the first time. Field studies in Natal have yielded additional information

on the habitat, phenology, fruit morphology and pollination biology of T. macranthum (Bak.) C.H. Wr.

Phalaenophily is reported for the second time in the South African Liliaceae. The relationship of T. macranthum

with the moth Diaphone ettmela (Cramer) — Noctuidae, Hadeniae, is discussed.

Resume

THURANTHOS: NOTES SUR LE STATUT GENERIQUE, MORPHOLOGIQUE, PHENOLOGIQUE

ET BIOLOGIQUE DE LA POLLIN/SATION

Le slat lit generique de Thuranthos C. H. Wr. est discute. Une clef separant les deux especes est fournie. Le

genre est signale du Natal pour la premiere fois. Des eludes sur le terrain au Natal ont fourni des informations

additionnelles sur T habitat, la phenologie , la morphologie du fruit et la biologie de la pollinisation du T. macrant-

hum (Bak.) C.H. Wr. La phalaenophilie est signalee pour la seconde fois dans les Lileaceae d'Afrique du Sud. La

relation dt T. macranthum avec Talene Diaphone eumela (Cramer)— Noctuidae, Hadeniae, est discutee.

INTRODUCTION

Thuranthos C.H. Wr. provides an example of the

difficulties and controversies encountered in the

delimitation of genera in the Liliaceae. Wright (1916a)

established Thuranthos to accommodate the species

Thuranthos macranthum (Bak.) C.H. Wr. which had

been anomalously referred to the genera Ornithogalum

L. and Drimia Jacq. respectively (Fig. I). A second

species T. nocturnale was described by R. A. Dyer

in 1964 (Fig. 2). The generic name Thuranthos is

derived from Ovpa, an opening, and avOo $, a flower,

in allusion to the interspaces between the lower parts

of the filaments (Wright, 1916b).

GENERIC STATUS OF THURANTHOS

In a recent study Jessop (1973) sinks Thuranthos

C.H. Wr., Urginea Steinh. and (Jrgineopsis Compton

under Drimia Jacq. This decision is clearly based

on the relationship between Urginea and Thuranthos

on the one hand and between Urginea and Urgineopsis

on the other. Field studies suggest, however, that the

two species of Thuranthos have very distinct features

separating them from all species of the other three

genera.

Like other authors (Wright 1916a, 1916b; Dyer,

1964) I consider the structure of the filaments of

Thuranthos to be an outstanding feature, which is of

particular significance with respect to pollination

biology. The structure of the filaments only becomes

visible once the perianth segments have reflexed and

for this reason is scarcely apparent in herbarium

specimens.

The filaments are clearly divisible into two

regions. Fig. 3.1 shows the basal parts which are

fl ittened and convex. This region resembles a paper

lantern with longitudinal slits, in the upper region

(Fig. 3.2) the filaments are mostly terete and straight

but often arch near the anthers (Fig. 3.3) and at the

point of connivence (Fig. 3.4). The basal region is a

drab yellow-green, like the perianth segments, whereas

the upper region is white. The green anthers produce

white lanceolate pollen grains (Fig. 3.5).

* Botanical Research Institute, Department of Agricultural

Technical Services, Private Bag X101, Pretoria.

mt'

Fig 1 — Thuranthos macranthum (Bak.) C.H. Wr., photograph

'from Bot. Mag. CXLII. t. 8680 (1916).

162 THURANTHOS: NOTES ON GENERIC STATUS, MORPHOLOGY, PHENOLOGY AND POLLINATION BIOLOGY

SPECIES OF THURANTHOS

Fig. 2. — Thuranthos nocturnale R. A. Dyer, photograph from

Flow. PI. Afr. 36. t. 1439 (1964).

Fig. 3. — Thuranthos maeranthum, showing open flower. 1, basal

region of androecium; 2, upper region of androecium;

3, arching base of upper filament region; 4, arching apex

of upper filament region; 5, pollen grain; 6, T/S near apex

of ovary, gland position indicated in corners, style position

shown in centre; 7, papillae of stigma.

None of the species of Urginea, Urgineopsis or

Drimia has such a complex filament structure. Until

detailed field studies on the Drimia — Thuranthos

relationship (sensu Jessop, 1973) have been under-

taken it seems best to maintain Thuranthos as a

genus.

1. Thuranthos maeranthum ( Bak .) C.H. Wr. in Kew

Bull. 1916: 213 (1916); C.H. Wr. in Bot. Mag.

CXLTI. t. 8680 (1916).

Ornithogalum maeranthum Bak. in J. Linn. Soc. (Bot).

13: 280 (1873). Types: Cape Province, Drege 2204, 3531.

Drimia macrantha (Bak.) Bak. in Bot. Jahrb. 15 (3): 7 (1892)

Urginea macrantha (Bak.) Phill. in Ann. S. Afr. Mus. 16: 305

(1917).

2. Thuranthos nocturnale R. A. Dyer in Flow. PI.

Afr. 36: t. 1439 (1964). Type: Cape Province,

Grootfontein, Acocks 18650 (PRE, holo!).

In the past these two species have been separated

on the shape of the leaves and capsules, the presence

or absence of pubescence on the stigma and the

length of the peduncle. The following key incorporates

additional characters observed during field studies.

Scape 50-140 cm high, peduncle 1-J- to 2 X longer than

inflorescence, not noticeably thickened at base; flowers

pendulous, pedicels of reflexed flowers arched, only

inner perianth segments tightly reflexed; stigma

glabrous; capsule (3, 4)4, 0-5,0 cm long, 1, 5-2,0 cm

broad, ovate oblong, twice as long as broad, upright on

vertical pedicel, surface smooth maeranthum

Scape 45-60 cm high, peduncle same length as inflores-

cence, noticeably thickened at base; flowers angled

20°-30° from rhachis with apex nearest rhachis, pedicels

of reflexed flowers almost horizontal, all perianth

segments rightly reflexed, stigma pubescent; capsule

3, 0-3, 5 cm long, 2, 5-3,0 cm broad, oblong, as long

as broad, upright on horizontal pedicel, surface

wrinkled nocturnale

BIOLOGY OF T. MACRANTHUM IN NATAL

POPULATIONS

Distribution

Two populations of T. maeranthum have been

found in Natal and seem to constitute the second

record for the Province. Ross (1972) cited Taylor

29750 (NH), from the Natal Midlands, as T. macran-

thum. I have not seen this specimen. The additional

Natal localities are Edendale, Pietermaritzburg (2931

CB) and Mkondeni (2931 CB).

Habitat

The Edendale population occurs in an annually

burnt vlei. This area holds surface water in shallow

depressions throughout the summer but gradually

dries out with the onset of winter by which time very

little vegetation is photosynthesizing. Plants were

found to grow on raised tussocks.

The Mkondeni population occurs along the edge

of a well-grassed donga (eroded gulley) which,

although drier than the Edendale population, generally

holds more water than the surrounding area.

The Edendale locality has a soil belonging to the

Rensburg series whereas the Mkondeni locality has a

soil of the Clovelly Series. These soils differ markedly

in PH value, base saturation and the sum total of

exchangeable cations (Van der Eyk, Macvicar &

De Villiers, 1969). From this information one could

conclude that this species is plastic as to its general

edaphic requirements.

C. H. STIRTON

163

Fig. 4. — Thuranthos macranthum, rhachis bearing fruits

1, erect pedicel; 2, upright deeply angled fruit. Note the

remains of the perianth segments on the right hand fruit.

The centre fruit is atypical in form.

Phenology

Scapes are produced during October and November.

The erect 50-140 cm scapes emerged about three

months before the leaves. Nineteen scapes emerged

in the Edendale locality during 1973, and over sixty-

five scapes were counted during 1974. This seems to

indicate a seasonal flowering periodicity and needs

further study. In both localities the scapes emerged in a

flush with no recorded stragglers appearing at a

later date. An average of 15 to 24 flowers were

produced per plant in both populations. Leaves

appeared during late January after the capsules had

dehisced and the scapes had died back.

Fruit morphology

Capsules were unknown to Jessop (1973). A short

description, however, did appear in Bot. Mag.

CXLII t. 8680 (1916): “Capsula longa, laevis, 3,5 cm

longa, 1 cm diametro, pedicillis suberectus instructa”.

Based on a population of 135 capsules, from two

populations, this description is amended as follows.

Capsule upright on erect pedicel (Fig. 4.1) deeply

3-angled (Fig. 4.2), (3.4) 4, 0-5,0 cm long, 1, 5-2,0

cm broad, narrowing from base to apex, base slightly

cordate to truncate, apex emarginate-truncate, more

than twice as long as broad, carpels with slight

groove down angle (Fig. 5.1), surface smooth (Fig.

5.2), green with a faint white bloom, turning brown

with age, basipetal tricarpellar dehiscence. Seeds

10-12 mm long, 6-8 mm wide, black.

Fig. 6. — Diaphone eumela (Cramer), moth and larval stages.

Fig. 5. — Fruit of Thuranthos macranthum. The fruit shows

feeding damage by the caterpillar Diaphone eumela

(Cramer) shown in the foreground. 1, slight groove down

angle; 2, smooth surface of capsule.

Pollination biology

Important features of the flower of T. macranthum

as shown below agree with those given by Faegri &

Van der Pijl (1971) as typical of phalaenophily (moth

pollination).

Nocturnal anthesis

Diurnal closure

Drab colour

Copious secretion of strongly scented nectar

Versatile anthers

Deeply divided perianth segments (reflexing

upwards)

Lattice like structure of basal region of androecium

and of reflexed perianth segments

Partly obscured nectar.

Field studies have shown that the Noctuid moth

Diaphone eumela (Cramer), Fig. 6, is closely

associated with T. macranthum. The relationship is

discussed after a short description of the floral

mechanism of T. macranthum.

1 64 THURANTHOS: NOTES ON GENERIC STATUS, MORPHOLOGY, PHENOLOGY AND POLLINATION BIOLOGY

Open flowers exude strongly perfumed nectar

which saturates the ambient atmosphere. The nectar is

produced by three glands, one each at a corner of

the triangular apex of the ovary (Fig. 3.6). A drop

of nectar is visible in Fig. 8.1. Note also the shiny

inner surface of the perianth segments which could

be due to some secretion (Fig. 8.2).

Flowers open only once. After closing in the

early morning the flowers dry out and by evening

they become mucilaginous owing to the breakdown

of tissue of the inner walls of the perianth segments.

No more than three flowers were ever seen open on

any one plant at the same time. The flowering period

of individuals extended from six to nine days

depending on the number of flowers produced per

inflorescence.

The moth Diaphone eumela (Cramer) belongs to

the family Noctuidae, sub-family Hadeniae (Fig. 6).

It lays its eggs on the youngest buds of the

inflorescence of T. macranthum usually at a time

when the first flower opens. The young larvae feed

on the flower buds. It was found that the first opened

flowers had produced sizeable fruits by the time the

larvae, which had undergone a number of instars,

had reached these fruits. Fruits and ovaries appeared

to be the main diet of grown larvae (Fig. 5). No more

than fifteen of these yellow and black caterpillars

were seen on any one plant, and were found on only

6 per cent of the total number of scapes produced in

both populations. On maturity the caterpillars crawl

down the scape and pupate beneath the soil, forming

mud cocoons. Laboratory reared caterpillars formed

chrysali without spinning a cocoon if no substrate

was provided, otherwise they pupated like the “wild

specimens”. Moths emerged from the chrysali after

a period of three weeks under laboratory conditions.

Fig. 7. — Thuranthos macranthum, flowers in the process of

opening. 1, outer perianth segments; 2, inner perianth

segments. Compare the position of the filaments with the

later stage shown in figure 8.

Flowers opened at dusk between 1730 and 1800

hours and closed at dawn between 400 and 500 hours.

Laboratory and field observations strongly suggest

that a circadian rhythym is involved. This pheno-

menon needs further study by physiologists.

Fig. 7 shows flowers in the process of opening.

The outer perianth segments are the first to spread

outwards and upwards (Fig. 7.1). They are followed

closely by the inner perianth segments (Fig. 7.2)

which curl rapidly and tightly inwards until the

tips almost touch the apex of the pedicel (Fig. 8.3).

The pedicel itself stiffens and arches so that it lies

above the horizontal. This movement forces the open

flower away from the rhachis (Fig. 3). In this aspect

T. macranthum differs markedly from T. nocturnale .

In the latter the pedicel is patent and only curves

downwards slightly at the tip. This forces the flower

well away from the rhachis and at about an angle of

20°-30& off the vertical (Fig. 2). It is worthwhile to

note that as far as the reflexure of perianth segments

is concerned both Fig. 1 and 2 are incorrect. These

paintings were probably made during the late

afternoon or early evening before the perianth

segments had completely reflexed. Mrs A. A. Ober-

meyer confirms this for Fig. 2. Coloured negatives of

the open flowers of the type specimen of T. nocturnale

(in PRE) show that all the perianth segments reflex

tightly towards the apex of the pedicel. This is not

shown in Flow. PI. Afr. 36. t. 1439 (1964).

p,G- 8. — Thuranthos macranthum, flowers open. 1, drop of

exuded nectar; 2, shiny undersurface of perianth segments;

3, tips of inner perianth segments reflexed tightly towards

the apex of the pedicel.

C. H. STIRTON

165

Diaphone eumela (Cramer) pollinates T. macranthum

as follows: The moth lands on the filament cage with

its head pointing upwards. The movement of its

feet as it attempts to maintain this position causes the

independant filaments to move. These filaments push

against the style at the point of connivence then slip

off (similar to pushing a pencil against a thin cylinder).

The moth usually slides down until it hangs beneath the

anther cage. This causes the moth’s abdomen to rub

against the anthers and sticky stigma. Some moths

were observed to land on the top of the perianth

segments, head downwards. The effect of their

movements resulted in the ready release of pollen.

In such cases very little pollen fell on the stigma as

only the apical part is receptive i.e. the half-circle

facing the ground. Note the stigmatic papillae in

Fig. 3.7.

Apart from D. eumela (Cramer) no other pollinators

were observed. Autogamy may occur but seems

unlikely in view of the small number of capsules

produced per plant. Moths were noted to visit most

open flowers, often returning to the same flower on

a number of occasions to feed on the nectar. There

is no record of pollination in T. nocturnale.

Stuckenberg (1975) informed me that D. eumela

had been recorded to have the following host plants:

Ornithogalum vir dm Lindl. (=0. ecklonii Schlechtd),*

Dipcadi viride (L.) Moench [=D. umbonatum (Bak.)

Bak.] and Albuca setosa Jacq. (=A. pachychlamus

Bak.). T. macranthum is a new host plant record.

Davidson (1975) has observed at least three further

Liliaceous genera on which the larvae of D. eumela

also feed. It would be most interesting to have further

information of the relationship, if any, of the moth

to the flowers of these various genera.

DISCUSSION

Judging by the existing literature and by my own

observations phalaenophily seems to be a rarely

recorded type of pollination in the South African

flora. Vogel (1954) records phalaenophilous flowers

for only one South African member of the Liliaceae,

namely a species of Dipcadi. This report constitutes,

to the best of my knowledge, the second such case.

The meagre knowledge of this phenomenon could

undoubtedly be due to the nocturnal behaviour of

both plants and insects, as opposed to the diurnal

closure of botanists and entomologists. Future

pollination studies in the South African flora could

conceivably be based on a critical assessment of

structural morphology with subsequent extrapolation

to the field.

ACKNOWLEDGMENTS

The author is indebted to the Transvaal Museum

for the loan of insect specimens; Dr L. Vari, Transvaal

Museum, for identifying the larvae and moth stages

of D. eumela ; Mr M. Scoble, Transvaal Museum,

Pretoria, and Dr B. Stuckenberg, Natal Museum,

Pietermaritzburg, for their interest and entomological

advice; Mrs A. A. Obermeyer and Dr O. A. Leistner,

Botanical Research Institute, Pretoria, for a critical

reading of the manuscript; Dr R. A. Dyer, Botanical

Research Institute, for interesting comments on the

genus Thuranthos', Miss R. Deakin, Rosebank,

Cape Town, for valuable assistance during field

studies; Miss M. Scott and Mrs Romanowski for

technical assistance.

UITTREKSEL

Die generiese status van Thuranthos C.H. Wr.

word behandel. 'n Sleutel vir die onderskeiding van die

twee species word verskaf. Vir die eerste keer word

aangeteken dat die genus in Natal voorkom. Veldwerk

in Natal bet addisionele inligting oor groeiplek, feno-

logie, vrugmorfologie en bestuiwingsbiologie van T.

macranthum (Bak.) C.H. Wr. aan die lig gebring.

Motbestuiwing word vir die tweede keer in die Suid-

Afrikaanse Liliaceae aangeteken. Die verwantskap

tussen T. macranthum en die mot Diaphone eumela

( Cramer ) — Noctuidae, Hadeniae, word bespreek.

REFERENCES

Davidson, L., 1975. Personal communication.

Dyer, R. A., 1964. Thuranthos nocturnale R. A. Dyer. Flow

PI. Afr. 36: t. 1439.

Faegri, K., van der Pul, L., 1971. The principals of pollination

ecology. 2nd Revised Ed. Oxford: Pergammon. 291 pp.

Jessop, J. P., 1973. Studies in the South African bulbous Liliaceae.

Ph.D. thesis, Rhodes University (Unpublished).

Stuckenberg, B., 1975. Personal communication.

Van Der Eyk, J. J., MacVicar, C. N., De Villiers, J. M.,

1969. Soils of the Tugela Basin. Natal Town and Regional

Planning Reports. No. 15. 263 pp.

Vogel, S., 1954. Bliitenbiologisclie Typen als Elemente der

Sippengliederung: DargesteUt aanhard der flora Siidafrikas.

Veb. Gustav Fischer Verlag: Jena. 338 pp.

Wright, C. H., 1916a. Diagnosis Africanae. LXIX. Bull.

Misc. Inf. 9: 233-234.

Wright, C. H., 1916b. Thuranthos macranthum. Bot. Mag.

CXLII : t. 8680.

167

Book Reviews

Cvcads of Southern Africa by Cynthia Giddy. Cape Town:

Purnell. 1974. Pp. 122, with numerous pencil drawings and 32

pages of colour reproductions. Pr.ce R 12,50.

Cynthia Giddy does not tell tlje reader what her aims or

objects were in writing this book. One-gains the impression that

it was an irresistible urge to share with everyone her wonderment

of the magic atmosphere created for her by the Cycads in

their native habitats, a feeling that she was living in the past

stretching into many millions of years. For those who know

their Cycads there is not a wealth of new information, never-

theless her presentation of the facts as they are known is gene-

rally pleasing. She can hardly be blamed if the book stimulates

the further collection of plants from the veld in spite of her

professed dislike of large cycads in suburban gardens and in

spite of the protective laws, which seem readily circumvented.

Having learnt from earlier publications that the habit of

growth and the characteristics of the male and female pones are

of prime importance in the classification of the species, she

has gone to infinite pains to gather this essential information

for each of the 28 recognized species of Encephalartos in

Southern Africa and one of Stangeria. She sought the assistance

of scientists and laymen and travelled long distances by car

and on foot in her quest for good coloured illustrations. Her

major personal contribution to the book are the 32 pages of

colour plates, illustrating the 28 species of Encephalartos , one

of Stangeria eriopus , one of hybrids and two of subjects of

general interest. On each of the pages there are up to six attrac-

tive photographs. As a reader though, I have an aversion to

being told to “note” the obvious, which is too frequently an

injunction in the captions to the coloured illustrations. The

colours are somewhat over-emphasized and one does not get the

same detail with colour reproductions as is possible with black

and white photographs. While on the topic of illustrations,

praise must be given for the pencil drawings contributed by

Barbara Jeppe. These consist mainl^ of diagrams, habit sketches

and leaf-shapes, which add appreciably to the interest of the

text. The text includes short chapters on reproduction, culti-

vation, propagation, toxicity and identification, which are all

informative, without being highly original. The author says of

her key that it attempts to group the species (which are unnum-

bered) according to their appearance and that it does not

necessarily reflect any direct relationship.

There is a confusion of ideas on p. 81 relative to hybrids.

It is stated that the distribution areas of E. villosus and E. umbelu-

ziensis do not overlap (from which one infers that there are no

natural hybrids), but she says that recent field work points

to the possibility of an intermediate species. For evidence of

this one is referred to Colour Plate 30, Figs. 5 and 6. By no

stretch of imagination could these figures be regarded as female

and male plants of a so-called “intermediate” species. No

information is given as to when and where the field work was

done and there is no guide for a follow up even if one had the

inclination to delve further.

The unenlightened reader might infer from the heading to

the Foreword (which incidentally follows the Acknowledge-

ments, Preface and References) that it was written by Prof.

Brian Rycroft and Harold Pearson. Brian Rycroft, in addition

to his post of Director of Kirstenbosch is the ‘Harold Pearson’

Professor of Botany at U.C.T. This chair of botany commemo-

rates the founder and first Director of Kirstenbosch. The

name Heenan is recorded as Heenen throughout the text and

the initials of R.A.D. are twice given prominently as R.H.D. —

curious oversights.

The publishers, Purnell of Cape Town, have placed their

stamp of quality on the book, in its style and in the materials

used. As soon as it was published in 1974 the book met with

an enthusiastic welcome from the public. In this it was a success-

ful venture.

It is common knowledge that in spite of the official regulations

for their protection in South Africa, in spite of the flood of

pious sentiment expressed by conservationists and others,

very many thousands of Cycads have been ruthlessly tom from

their ancient strongholds to satisfy the avarice of man. The

plundered species can never regain their magnificent pristine

glory. Both botanically and aesthetically the situation is a

tragedy.

R. A. Dyer

Handbook of Vegetation Science. Chief ed. R. Tuxen.

Vegetation and Environment. Ed. B. R. strain & W. D. Bil-

lings. The Hague : W. Junk. Vol. 6. 1974. Pp. VIII & 194, 5

photographs, 30 figures, 9 tables. Price R15.

This small volume presents a concise but sound review of the

nature of the interactions between vegetation and environment,

and it outlines recommendations for their analyses. The book

contains nine short chapters written by thirteen authors. Twelve

of these authors are American, seven being based in North

Carolina. This strategy in the choice of authors has resulted in a

particularly well-integrated text.

In the first chapter, Billings presents an excellent account of

the concept of environment and the ways of measuring it,

discussing topics like tolerance, ecotypic adaptations and

acclimatization of plants to environmental components. He

also explains the environment as a holocoenotic system and

shows some appalling results of trigger factors starting virtually

irreversible, destructive developments in some ecosystems.

Then there are chapters on ecological niche and on the possibi-

lities of mathematically describing plant-environment relation-

ships. Muller summarizes the published results of his and other

studies on the role of allelopathy in the environmental complex.

J. T. Scott tries to define the interacting system of vegetation

and environment in such a way that it will allow a decisive test

of the continuum and community-type hypotheses. This

excercise is, in my view, superfluous: it does not contribute

significantly to the general theme of the book.

Mooney contributes an interesting discussion on plant forms

in relation to environment. He states that for any given habitat

condition the dominant plant form represents the optimal

form-behavioural strategy for carbon gain. This plant type

cannot utilize all resources, however, and the result is a resource

division allowing for other plant types in the area. Mooney

also briefly mentions the IBP studies on convergent evolution

in two pairs of homoclimatic areas in North and South America,

comparing Californian chaparral with Chilean matorral, and

the North American with the Argentinian desert area.

The book concludes with two chapters providing guidance

for further research, one dealing with the modelling of photo-

synthesis of plant stands, and the other discussing the experimen-

tal analysis of ecosystems.

Author and subject indices increase the usefulness of this

valuable little volume.

M. J. A. Werger

Multivariate Analysis in Vegetation Research by L.

Orloci. The Hague: Dr W. Junk B.V. 1975. pp. ix + 276

(paperbound). Appendix of glossary and listings of 20 basic

computer programs. Price Dutch Guilders 50.

Ldszlo Orloci, the well-known quantitative ecologist from

Canada’s University of Western Ontario, in a short book of

168 text pages presents material which he has used in a graduate

plant ecology course. He proposes that his book be used in

classes with an interest in multivariate data analysis and also

by research workers interested in broadening their experience

with data analysis. The book’s objectives, firstly, to provide

an introduction to the concepts and procedures that need be

known before application of multivariate analysis and, secondly,

to serve as a source for worked examples and useful computer

routines, are successfully reached in a clearly-understandable

fashion. In each of the six chapters the mathematical theory

of the subject is given and is followed by a small worked

example. Unusually clear listings of computer programs in the

basic language and results of computations with test data

in the 100-page appendix round off most. methods he describes

and makes it possible for anyone with access to a computer

to be able to use his methods of analysis.

South African ecologists, including those with quantitative

as well as Braun-Blanquet loyalties, will benefit from reading

the short, objectively-written section on sampling considerations.

He coins the useful term, preferential sampling, to describe

the choice of ‘typical’ stands of vegetation and sampling from

‘preferred’ sampling sites, as usually practised by Braun-

Blanquet adherents. He points out that such data will not be

suitable for statistical analysis but that systematic, random or

restricted-random sampling methods may be used to gather

data for later statistical analysis. The sampling method depends

on the problem being investigated, the variables to be measured,

the mathematical model to be used and the intended method

of data analysis. These considerations are clearly discussed and

useful guide-lines are laid down to assist an ecologist with the

design of his sampling strategy.

In the first chapter, he also describes a procedure which

enables one to calculate, from a pilot sample, the lowest number

of species which need be recorded in the main sample to obtain

a stated minimum level of distortion. The same procedure

calculates the sampling effort involved to obtain this level of

distortion: This technique could have wide application in

South Africa, where the few quantitative ecologists must use

every possible aid to enable them to complete the tasks assigned

to them within a reasonable time.

The second chapter describes and evaluates various resem-

blance functions, which generate the input to ordination and

classification methods, described in chapters three and four,

respectively. Four different goals, for which ordination methods

168

have been put in vegetation studies, are listed. One of these,

summarization, is the most usual goal of ordinations undertaken

in South Africa. For summarization, component analysis is

efficient and indicated, according to Orloci, if all sample points

fall in a single linear cluster. The well-known Bray & Curtis

method is considered by Orloci to be the least recommendable

multidimensional scaling method, even after the recent attempts

to restore it to the role which it once played in ecology. While

pointing out possible revisions to and improvements on the

mathematics, he draws attention to three roots of the problem,

namely, that the Bray & Curtis resemblance function often

lacks a consistent scale, axes are potentially non-intersecting

and oblique (correlated) and a method which produces scatter-

diagrams without taking into account the potential obliqueness

of the ordination axes is dubious.

The need for another revision and expansion of Greig-

Smith’s poineering Quantitative Plant Ecology has been felt

for some years and Orloci’s book succeeds in covering the

mathematical aspects of quantitative ecology most adequately.

Although the book draws heavily on the author’s past research

and includes some of his previously unpublished techniques,

the important findings of other quantitative ecologists are

quoted and discussed in the text. This book would be difficult

for a beginner in this field to grasp without guidance, as much

is either assumed or mentioned, in passing, together with a

literature reference. It is highly recommended as one of a few

class text-books for a post-graduate course in quantitative

ecology and as a source book for researchers in the field. The

need for an integrated, advanced text on ecological concepts

and methods, specifically written for South African students

and quoting the results of South African research grows with

advances in ecology such as those described in this and other

recent ecological texts.

J. W. Morris

GUIDE FOR AUTHORS

GIDS VIR SKRYWERS

GENERAL

Bothalia is a medium for the publication of botanical

papers dealing with the flora and vegetation of Southern

Africa. Papers submitted for publication in Bothalia should

conform to the general style and layout of recent issues of

the journal (from Vol. 1 1 onwards) and may be written in

either English or Afrikaans.

TEXT

Manuscripts should be typed, double-spaced on one side

of uniformly-sized A4 paper having at least a margin of 3 cm

all round. Latin names of plants should be underlined to

indicate italics. All other marking of the copy should be left

to the editor. Metric units are to be used throughout. Manus-

cripts should be submitted in duplicate to the Editor, Bothalia,

Private Bag X101, Pretoria.

ABSTRACT

A short abstract of 100-200 words preferably in both

English and Afrikaans should be provided. In the abstract

the names of new species and new combinations should not

be underlined.

FIGURES

Black and white drawings, including graphs, should be in

jet-black Indian ink preferably on bristol board. Lines should

be bold enough to stand reduction. Indicate the desired lettering

lightly in pencil: the printer will insert the final lettering. If

authors prefer to do their own lettering, then use some printing

device such as stencilling, letraset, etc. It is recommended

that drawings should be twice the size of the final reduction.

Photographs submitted should be of good quality, glossy,

sharp and of moderate, but not excessive contrast. Photograph

mosaics should be composed by the authors themselves: the

component photographs should be mounted neatly on a

white card base leaving a narrow gap between each print;

number the prints using some printing device.

Figures should be planned to fit, after reduction, into

a width of 8 cm, 11 cm or 17 cm with a maximum vertical

length of 24 cm.

The number of each figure and the author’s name should

be written on the back of the figure using a soft pencil.

Captions for figures should be collected together and typed

on a separate page headed Captions for Figures. A copy of

each caption should be attached to the base of each figure.

Do not underline plant names in captions — only collectors’

names and numbers.

Authors should indicate in pencil in the text where they

would like their illustrations to appear.

TABLES

Tables should be set out on separate sheets and numbered

in Arabic numerals.

CITATIONS OF SPECIMENS

In citing specimens the grid reference system should be

used (Technical Note: Gen. 4, 4c). Provinces/countries should

be cited in the following order: S.W. Africa, Botswana,

Transvaal, Orange Free State, Swaziland, Natal, Lesotho and

the Cape. Grid references should be cited in numerical sequence.

Locality records for specimens should preferably be given to

within a quarter-degree square. Records from the same one-

degree square are given in alphabetical order i.e. ( — AC)

precedes ( — AD), etc. Records from the same quarter-degree

square are arranged alphabetically according to the collectors’

names; the quarter degree references must be repeated for each

specimen cited. The following example will explain the proce-

dure.

Natal. — 2731 (Louwsburg): 16 km E. of Nongoma (-DD),

Pelser 354; near Dwarsrand, Van der Merwe 4789. 2829

(Harrismith): near Groothoek (-AB), Smith 234; Koffiefontein

(-AB), Taylor 720; Cathedral Peak Forest Station (-CC),

Marriott 74; Wilgerfontein, Roux 426. Grid ref. unknown:

Sterkstroom, Strydom 12.

Records from outside Southern Africa should be cited

from north to south i.e. preceding those from Southern Africa.

The abbreviation “distr.” should be added to all district

names, e.g.

Kenya. — Nairobi distr.: Nairobi plains beyond race course,

Napier 845.

ALGEMEEN

Bothalia is ’n medium vir die publikasie van plantkundige

artikels wat handel oor die flora van Suidelike Afrika. Artikels

wat voorgele word vir publikasie in Bothalia behoort ooreen

te stem met die algemene styl en rangskikking van onlangse

uitgawes van die tydskrif (vanaf Vol. 11). Dit mag in Engels

of in Afrikaans geskryf word.

TEKS

Manuskripte moet getik wees in dubbelspasiering slegs

op een kant van ewegroot A4 papier, met reg rondom ’n

rand van minstens 3 cm breed. Latynse name van plante moet

onderstreep word om aan te dui dat dit kursief gedruk moet

word. Alle ander merke moet aan die redakteur oorgelaat

word. Metrieke eenhede moet deurgaans gebruik word.

Manuskripte moet in tweevoud ingedien word by die Redakteur,

Bothalia, Privaatsak X101, Pretoria.

UITTREKSEL

’n Kort uittreksel van 100-200 woorde moet voorsien

word, verkieslik beide in Engels en Afrikaans. In die uittreksel

moet die name van nuwe soorte en nuwe kombinasies nie

onderstreep word nie.

AFBEELDINGS

Wit en swart tekeninge, insluitende grafieke, moet met

pikswart Indiese ink geteken word, verkieslik op “bristol

board”. Lyne moet dik genoeg wees om verklein te kan word.

Dui die verlangde byskrifte liggies in potlood aan: die drukker

sal die uiteindelike byskrifte invoeg. Indien skrywers verkies

om hulle eie byskrifte te maak, gebruik dan een of ander

hulpmiddel soos letraset of ’n sjabloon. Dit is wenslik dat

tekeninge tweemaal so groot as die uiteindelik verkleining sal

wees.

Foto’s wat ingedien word, moet van hoe kwaliteit wees

glansend, skerp en van matige maar nie oordrewe kontras.

Fotomosai'eke moet deur die skrywer self saamgestel word:

die afsonderlike foto’s moet netjies monteer word op ’n stuk

wit karton met ’n smal strokie tussen die foto’s; nommer die

foto’s met behulp van een of ander druk-hulpmiddel.

Afbeeldings moet so beplan word dat hulle na verkleining

sal pas in ’n breedte van 8 cm, 1 1 cm of 17 cm met ’n maksimum

vertikale lengte van 24 cm.

Die nommer van elke afbeelding sowel as die skrywer

se naam moet op die rugkant van die afbeelding geskryf word

met ’n sagte potlood.

Onderskrifte vir afbeeldings moet bymekaar getik word

op ’n afsonderlike bladsy met die opskrif Onderskrifte vir

Afbeeldings. ’n Afskrif van elke onderskrif moet aan die

onderkant van elke afbeelding vasgeheg word. Moenie plant-

name in onderskrifte onderstreep nie, slegs versamelaarsname

en-nommers.

Skrywers moet met potlood in die teks aandui waar hulle

graag hulle afbeeldings wil he.

TABELLE

Tabelle moet op afsonderlike velle papier kom en genom-

mer word met Arabiese nommers.

SITERING VAN EKSEMPLARE

Wanneer eksemplare siteer word, moet die ruitverwysing-

stelsel gebruik word (Tegniese Nota: Gen. 4, 4c). Provinsies/

lande moet in die volgende volgorde siteer word: Suidwes-

Afrika. Botswana, Transvaal, Oranje-Vrystaat, Swaziland,

Natal, Lesotho en die Kaapprovinsie. Ruitverwysings moet

in numeriese volgorde siteer word. Lokaliteitsrekords vir

eksemplare moet verkieslik tot binne kwartgraadvierkante

gegee word. Rekords uit dieselfde eengraadvierkant word in

alfabetiese volgorde aangebied, nl. (-AC) kom voor (-AD),

ens. Rekords uit dieselfde kwartgraadvierkant word alfabeties

gerangskik volgens die versamelaars se name, en die kwart-

graadverwysings moet herhaal word vir elke eksemplaar wat

siteer word. Die volgende voorbeeld sal die metode verduidelik:

Natal. — 2731 (Louwsburg): 16 km O. van Nongoma

(-DD), Pelser 354; naby Dwarsrand, Van der Merwe 4789,

2829 (Harrismith): naby Groothoek (-AB), Smith 234; Koffie-

fontein (-AB), Taylor 720; Cathedral Peak Bosboustasie

(-CC), Marriott 74; Wilgerfontein, Roux 426; Ruitverwysing

onbekend : Sterkstroom, Strydom 12.

Rekords van buite Suidelike Afrika moet siteer word

van noord na suid, d.w.s. dit gaan die van Suidelike Afrika

vooraf. Die afkorting “distr.” behoort by alle distriksname

gevoeg te word, bv.

Kenya. — Nairobi-distr. : Nairobivlakte anderkant die ren-

baan, Napier 845.

REFERENCES

References in the text should be cited as follows: “Jones

(1955) stated. . or . . (Smith, 1956)” when giving

a reference simply as authority for a statement. The list of

references at the end of the article should be arranged alpha-

betically and the literature abbreviations used should conform

to the list of Literature Abreviations (Technical Note: Tax.

6/1) issued by the Botanical Research Institute, thus:

Hutchinson, J., 1946. A botanist in Southern Africa. London:

Gawthorn.

Morris, J. W., 1969. An ordination of the vegetation of

Ntshongweni, Natal. Bothalia 10: 89-120.

If, as in many taxonomic papers, periodicals or books

are mentioned in the text, usually in the species synopsis, they

should be cited as in the following examples: Gilg & Ben. in

Bot. Jahrb. 53: 240 (1915) and Burtt Davy, FI. Transv. 1:

122 (1926).

REPRINTS

Authors receive 75 reprints gratis. If there is more than

one author, this number will have to be shared between or

among them.

VERWYSINGS

Verwysings in die teks moet as volg siteer word: “Jones

(1955) beweer . . of “. . . (Smith, 1956)” wanneer ’n

verwysing slegs as outoriteit vir ’n stelling gegee word. Die

verwysingslys aan die einde van die artikel moet alfabeties

gerangskik wees en die literatuurafkortings wat gebruik word,

moet in ooreenstemming wees met die lys van Literatuur-

afkortings (Tegniese Nota: Tax. 6/1) wat uitgegee is deur die

Navorsingsinstituut vir Plantkunde, as volg:

Hutchinson, J., 1946. A botanist in Southern Africa. London:

Gawthorn.

Morris, J. W., 1969. An ordination of the vegetation of

Ntshongweni, Natal. Bothalia 10: 89-120.

Wanneer, soos in baie taksonomiese artikels die geval

is, tydskrifte of boeke in die teks genoem word, gewoonlik

in die soortsinopsis, behoort hulle siteer te word soos in die

volgende voorbeelde: Gilg & Ben. in Bot. Jahrb. 53: 240

(1915) en Burtt Davy, FI. Transv. 1 : 122 (1926).

HERDRUKKE

Skrywers ontvang 75 herdrukke gratis. Wanneer daar

meer as een skrywer is, sal hierdie aantal tussen hulle verdeel

moet word.

'

PUBLIKASIES VAN DIE NAVORSINGSINSTITUUT VIR PLANTKUNDE

Beskikbaar van die Afdeling Landbou-inligting, Departement van Landbou-tegniese Dienste, Privaatsak X144, Pretoria, Republiek

van Suid-Afrika.

FLORA VAN SUIDELIKE AFRIKA

’n Taksonomiese behandeling van die flora van die Republiek van Suid-Afrika, Lesotho, Swaziland en Suidwes

Afrika.

Sal bestaan uit 33 volumes, nie in numeriese volgorde nie.

Reeds beskikbaar:

Vol. 1 (1966). Prys Rl,75. Oorsee: R2,20. Pos vry.

Vol. 13 (1970). Prys RIO. Oorsee: R12. Pos vry.

Vol. 26 (1963). Prys R.4,60. Oorsee: R5,75. Pos vry.

DIE BLOMPLANTE VAN AFRIKA

Hierdie publikasie word uitgegee as ’n geillustreerde reeks, baie na die aard van Curtis se “Botanical Magazine”. Die

doel van die werk is om die skoonheid en variasie van vorm van die flora van Afrika aan die leser bekend te stel, om

belangstelling in die studie en kweek van die inheemse plante op te wek, en om plantkunde in die algemeen te bevorder.

Die meeste van die illustrasies word deur kunstenaars van die Navorsingsinstituut vir Plantkunde gemaak, dog die

redakteur verwelkom geskikte bydraes van ’n wetenskaplike en kunsstandaard afkomstig van verwante inrigtings

Onder huidige omstandighede word twee dele van die werk gelyktydig gepubliseer, maar met onreelmatige tussenpose;

elke deel bevat 10 kleurplate. Intekengeld bedra R1 ,50 per deel: 4 dele per band. Vanaf Band 27 is die prys per band

in linne gebind RIO, 00; in moroccoleer gebind R14,00.

BOTHALIA

Bothalia is ’n medium vir die publikasie van plantkundige artikels oor die flora en plantegroei van Suidelike Afrika.

Een of twee dele van die tydskrif word jaarliks gepubliseer.

Die volgende dele is beskikbaar:

Vol. 3 Deel 1 Uit druk

2 1937 75c

3 1938 75c

4 1939 75c

Vol. 4 Deel 1 1941 75c

2 1942 75c

3 1948 75c

4 1948 75c

Vol. 5 1950 R3

Vol. 6 Deel 1 1951 Rl,50

2 1954 R2, 50

3 1956 R2

4 1957 R2

Vol. 7 Deel 1 1958 R2

2 1960 R3

3 1961 R3

4 1962 R3

Vol. 8 Deel 1 1962 R3

2 1964 R3

3 1965 R3

4 1965 R3

Supplement

Vol. 9 Deel 1 1966 R3

2 1967 R3

3 & 4 1969 R6

Vol. 10 Deel 1 1969 R3

2 1971 R3

3 1971 R3

4 1972 R3

Vol. 11 No. 1 & 2 1973 R6

3 1974 R3

4 1975 R3

MEMOIRS VAN DIE BOTANIESE OPNAME VAN SUID-AFRIKA

Die memoirs is individuele verhandelings, gewoonlik ekologies van aard, maar soms handel dit oor taksonomiese

of ekonomiese-plantkundige onderwerpe. Veertig nommers is reeds gepubliseer waarvan sommige uit druk is.

BOTHALIA

Vol. 12, No. 1

APRIL 1976

CONTENTS -INHOUD

Page

Bladsy

1. The South African species of Hemizygia (Lamiaceae). L. E. Codd 1

2. The genus Syncolostemon (Lamiaceae). L. E. Codd 21

3. Studies in the Ericoideae. I. The genera Eremia and Eremiella. E. G. H. Oliver 29

4. Studies in the Ericoideae. II. The new genus Stokoeanthus. E. G. H. Oliver 49

5. Notes on African plants

Asclepiadaceae. R. A. Dyer 53

Blechnaceae. J. C. Scheepers and P. Vorster 57

Ericaceae. E. G. H. Oliver 57

Fabaceae. J. H. Ross 59

Liliaceae. J. P. Jessop; A. A. Obermeyer; D. S. Hardy 60

Melastomataceae. J. H. Ross 62

Mesembryanthemaceae. H. R. Tolken and J. P. Jessop 63

6. A procedure for standardizing comparative leaf anatomy in the Poaceae. I. The leaf blade as viewed

in transverse section. R. P. Ellis 65

7. Studies in the Hypoxidaceae. I. Vegetative morphology and anatomy. M. F. Thompson Ill

8. Notes on Veronaea including V. compacta sp. nov. M. C. Papendorf 119

9. The soil mycoflora of an Acacia karroo Community in the Western Transvaal. M. C. Papendorf 123

10. The mycoflora of wheat field debris, Part II. W. J. Jooste 129

11. Hans Joachim Schlieben, collector extraordinary. O. A. Leistner 133

12. A phytosociological classification of the Nylsvley Nature Reserve. B. J. Coetzee, F. van der

Meulen, S. Zwanziger, P. Gonsalves and P. Weisser 137

13. Thuranthos: notes on generic status, morphology, phenology and pollination biology. C. H. Stirton 161

Book Reviews 167

PRICE R5

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