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Bothalia

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

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’n Taksonomiese verhandeling oor die flora van die Republiek

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BOTHALIA

’N TYDSKRIF VIR PLANTKUNDIGE NAVORSING

A JOURNAL OF BOTANICAL RESEARCH

Volume 18,2

Editor/ Redakteur: O.A. Leistner

Assisted by B.A. Momberg

D.F. Cutler

B. de Winter

P.H. Raven

J.P. Rourke

M.J. Werger

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Editorial Committee / Redaksiekomitee

B. de Winter

D.J.B. Killick

O.A. Leistner

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M.C. Rutherford

J.C. Scheepers

ISSN 0006 8241

Issued by the Botanical Research Institute, Department of Agriculture and Water Supply, Private Bag X101, Pretoria 0001,

South Africa

Uitgegee deur die Navorsingsinstituut vir Plantkunde, Departement van Landbou en Watervoorsiening, Privaatsak X101,

Pretoria 0001, Suid-Afrika

1988

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CONTENTS - INHOUD

Volume 18,2

1. A revision of the genus Prionanthium (Poaceae: Arundineae). GERR1T DAV1DSE

2. Studies in the genus Riccia (Marchantiales) from southern Africa. 10. Two new white-scaled species of

the group ‘Squamatae’: R. argenteolimbata and R. albornata. O.H. VOLK, S.M. PEROLD and

T. BORNEFELD ,-5

3. Taxonomy and leaf anatomy of the genus Ehrharta (Poaceae) in southern Africa: the Dura group. G.E.

GIBBS RUSSELL and R.P. ELLIS 165

4. The Oxygonum dregeanum complex (Polygonaceae). G. GERMISHUIZEN 2 73

5. Notes on African plants:

Brassicaceae. Heliophila cornellsbergia, a new species from the Richtersveld. B.J. PIENAAR and

A. NICHOLAS 183

Ericaceae. Coilostigma zeyherianum — a correction. E.G.H. OLIVER

Geraniaceae. The correct author citation for Pelargonium section Otidia. P. VORSTER

Orchidaceae. Additions to the synonymy of Eulophia schweinfurthii. A. V. HALL jg^

Polygonaceae. Reinstatement of Oxygonum acetosella Welw. G. GERMISHUIZEN jg7

6. Fusarium tricinctum (Fungi: Hyphomycetes) in South Africa- morphology and pathogenicity.

SANDRA C. LAMPRECHT, W.F.O. MARASAS, P.S. VAN WYK and P.S. KNOX-DAVIES .... 189

7. Leaf anatomy of the South African Danthonieae (Poaceae). XVII. The genus Chaetobromus . R.P. ELLIS j 95

8. Flora of the Zuurberg National Park. 1 . Characterization of major vegetation units. B-E. VAN WYK,

P.A. NOVELLIE and C.M. VAN WYK 211

9. Flora of the Zuurberg National Park. 2. An annotated checklist of ferns and seed plants. B-E. VAN

WYK, C.M. VAN WYK and P.A. NOVELLIE 221

10. A synopsis of the plant communities of Swartboschkloof, Jonkershoek, Cape Province. D.J. McDONALD 233

11. A checklist of the flowering plants and ferns of Swartboschkloof, Jonkershoek, Cape Province. D.J.

McDONALD and M. MORLEY 261

12. Threatened plants of the eastern Cape: a synthesis of collection records. D.A. EVERARD 271

13. Description of a proteoid-restioid stand in Mesic Mountain Fynbos of the south-western Cape and some

aspects of its ecology. G. DAVIS 279

14. Miscellaneous notes:

The plant number scale — an improved method of cover estimation using variable-sized belt tran-

sects. R.H. WESTFALL and M.D.PANAGOS 289

15. New taxa, new records and name changes for southern African plants. G.E. GIBBS RUSSELL, W.G.

WELMAN, G. GERMISHUIZEN, E.'RETIEF, B.J. PIENAAR, C. REID, L. FISH, J. VAN ROOY,

C.M. VAN WYK, E.KALAKE and STAFF 293

16. Annual report of the Botanical Research Institute 1987/88 305

17. Obituary: Hermann Merxmuller (1920— 1988). JURKE GRAU (Translation: O.A. Leistner) 325

18. Book reviews 329

Bothalia 18,2: 143-153 (1988)

A revision of the genus Prionanthium (Poaceae : Arundineae)

GERRIT DAVIDSE*

Keywords: Arundineae, Cape, chromosome numbers, conservation, Poaceae, Prionanthium, taxonomic revision

ABSTRACT

Prionanthium is revised and three species are recognized. All three species have a chromosome number of n =7.

The genus is unusual in having multicellular, secretory glands on the glumes and a wide range of inflorescence

types. Prionanthium is considered to be arundinoid on the basis of its leaf anatomy, ciliate ligule, spikelet mor-

phology and, less strongly, chromosome number. Its chromosome number, multicellular glands, two florets per

spikelet, type of palea and rachilla extension relate it more closely to Pentaschistis than to any other extant

arundinoid genus. A fully illustrated taxonomic account is presented.

UITTREKSEL

Prionanthium word hersien en drie spesies word onderskei. A1 drie spesies het ’n chromosoomgetal van n =7.

Die genus is buitengewoon omdat dit meersellige afskeidingskliere op die glumas het en verskillende tipes bloei-

wyses het. Op grond van die blaaranatomie, gesilieerde tongetjie, blompakkiemorfologie en, in ’n mindere mate,

chromosoomgetal behoort Prionanthium tot die Arundineae. Die chromosoomgetal, meersellige kliere, twee

blommetjies per blompakkie, tipe palea en ragillaverlenging dui eerder op ’n verwantskap met Pentaschistis as

met enige ander bestaande genus in die tribus. ’n Volledige taksonomiese beskrywing word gegee.

INTRODUCTION

Prionanthium , a genus of three annual species, found

only in the Cape Province, is one of the rarest grass gene-

ra of southern Africa and one of the species has been

listed as endangered (Hall & Veldhuis 1985). The total

number of separate populations known is 14. While col-

lecting cytological specimens of grasses from the Cape,

we were able to locate populations of all three species

and this stimulated the following taxonomic revision.

Ellis (in prep.) will report in detail on the leaf blade

anatomy.

HISTORICAL review

The genus Prionanthium was first described by Des-

vaux (1831), who was unable to relate it to any other

genus known to him. He named the only species/*, rigi-

dum and gave its distribution as ‘India Oriental’, (India

and the East Indies). This was clearly an error, for the

illustration that he provided clearly indicates that P. ri-

gidum is the Cape species now known as P. dentatum.

Prionanthium was undoubtedly based on Thunberg’s

collection from the Cape, the only collection known of

the species until 1975, although this collection was not

specifically cited by Desvaux. However, Thunberg’s

collection had already been described by Linnaeus the

Younger (1781) as Phalaris dentata so that the correct

name for the type species, as pointed out by Henrard

(1941), is Prionanthium dentatum (L. f.) Henrard.

Nees (Lindley 1836) redescribed the genus as Prio-

nachne, basing it on the second known species, P. eck-

lonii, and later (Nees 1841) illegitimately renamed Prio-

nachne as Chondrolaena .

* Missouri Botanical Garden, P.O. Box 299, St Louis, Missouri

63166, USA.

MS. received: 1988.03.30.

The third species, P. pholiuroides , was described by

Stapf (1899).

For more than two centuries, the genus was asso-

ciated with a mixture of genera (often recognized as

subtribes (cf. Pilger 1954; Zotov 1963)) that would be

termed avenoid and arundinoid in modern terms (cf.

Clayton & Renvoize 1986). Following Hubbard’s (1948)

formal recognition of the tribe Danthonieae, Chippindall

(1955) was the first to explicitly and exclusively associate

Prionanthium with arundinoid genera in the modem

sense of Clayton & Renvoize (1986). This placement has

been followed by De Wet (1956), Brown (1977), Ren-

voize (1981, 1986), Clayton & Renvoize (1986) and

Watson et al. (1 986).

MORPHOLOGY

There are several morphological features which merit

discussion beyond that provided in the formal descrip-

tions of the taxonomic section.

Multicellular, presumably secretory, glands occur on

the glumes of all three species. They are of two kinds:

stalked in P. dentation (Figures 1A, B; 2C) and P. eck-

lonii (Figure 1C, D) and sessile in P. pholiuroides (Figures

IE, F, H; 2D). The stalked glands are approximately

cylindrical in shape with sloping bases. They are com-

posed of 16 rows of cells in circumference. The apex is

rounded with a shallow, basin-like depression in the

centre surrounded by a low, heavily cutinized lip. The

apices of the glands illustrated for P. ecklonii (Figure

ID) have apparently been slightly distorted through

shrinkage. At the bottom of the basin-like depression

is a rectangular grid formed by the raised, rigid, anti-

clinal adjacent cell walls of small, square cells (Figure

1H). The stalks of the glands tend to be longer toward

the apex of the glume in both P. dentatum (Figure 1A)

and/*, ecklonii (Figure 1C).

144

Bothalia 18,2 (1988)

FIGURE 1. — Multicellular, secretory glands of Prionanthium. A-B, Prionanthium dentatum, Davidse

33394, on keel of glume: A, inner view of glume with stalked glands along the keel, X 20; B, stalked

glands, X 100. C-D, P. ecklonii, Davidse 34018 : C, inner view of glume with stalked glands along

the keel, X 20; D, stalked glands, X 100. E-H, P. pholiuroides : E-F, H, sessile glands on keel of

glume, Davidse 33983 ; E, inner view of glume, X 20; F, sessile glands, X 100; G, Davidse 34053,

glandless glume, keel with only prickles, X 100; H, close-up of sessile gland from Figure IF, X 360.

Bothalia 18,2 (1988)

145

FIGURE 2. — Morphology of Prionanthium species. A, habit

of P. ecklonti, Davidse 34018. B-D, inner view of glumes:

B, P. ecklonti, Davidse 34018\ C, P. dentatum, Davidse

33394 ; D, P. pholiuroides, Duthie sn.

The sessile glands of P. pholiuroides are basically

elliptical in outline to nearly circular in the smaller

ones (Figure IF, H). The cutinized Up arises directly

from the surface of the glume and is more massive than

in P. dentatum and P. ecklonti. The central depression

of the gland has the same type of grid as the other two

species (Figure IF, H).

The glands occur primarily along the midnerve of

both glumes, but they are completely absent in some

plants of three populations of P. pholiuroides (Struis-

baai, Davidse 34048, 34053 (Figure 1G); Stellenbosch

Flats, Duthie 1 76 7; Gordons Bay, Parker 3745). In one

other population ( Duthie s.n. in PRE-39169) of this

species, they may occur on some of the lateral nerves

in addition to the midnerve (Figure 2D).

Multicellular, secretory glands are rare in the Poaceae

as a whole, but they occur in about half of the species of

Pentaschistis, an arundinoid genus centred primarily in

South Africa. In the glandular Pentaschistis species, mul-

ticellular, secretory glands occur on the leaves and in-

florescence branches as well as on the glumes. Further-

more, in addition to stalked and sessile, multicellular

glands, sunken glands also occur, for example in the leaf

blade of P. angustifolia (Nees) Stapf and P. setifolia

(Thunb.) McClean (Ri\ Ellis pers. comm.).

For comparison, glands of two species of Pentaschistis

were examined in detail. In P. airoides (Nees) Stapf, the

glands on the inflorescence branches are fundamentally

similar to those of Prionanthium pholiuroides in being

sessile with a prominent, heavily cutinized Up, surround-

ing a central depression with a reticulate grid (Figure

3A). In the marginal, sessile leaf glands of P. aspera, the

internal structure of the depression is more elaborate.

The longitudinal cell walls of the grid in the central de-

pression have developed to a size approximately half as

large as the lip, and so form prominent ribs (Figure 3B).

Evidently, Pentaschistis has a more diverse array of glands,

although at least one kind is fundamentally similar to

those of Prionanthium.

Multicellular glands also occur in many species of

Eragrostis where they are always sessile, in the Clavelli-

gera group of Panicum, where they are prominently

stalked, and in Sporobolus heterolepis (A. Gray) A. Gray,

where they are raised bands on the pedicels (Bessey

1884). In some Eragrostis species the basic structure is

quite similar to that of Prionanthium (Nicora 1941),

whereas those of Panicum are quite different with long,

slender stalks supporting a spherical cluster of paren-

chyma cells and have been termed multicellular glandu-

lar macrohairs (Kabuye & Wood 1969).

The function of the glands in Prionanthium is not

known. In some freshly collected specimens of P. pholi-

uroides the glands were glistening with a thin layer of

moisture which I assume to indicate active secretion.

In very glandular species of Pentaschistis, such a &P. aspera

(Thunb.) Stapf, P. angulata (Nees) Adamson,/*, angusti-

folia (Nees) Stapf and others, the plants may be sticky

and/or produce an unpleasant smell. The volatile sub-

stance that is produced is not known. I did not detect

any kind of odour in any species of Prionanthium. In

Pentaschistis, at least in those species that produce it

146

Bothalia 18,2 (1988)

FIGURE 3. — Multicellular, secre-

tory glands of Pentaschistis .

A, Pentaschistis airoides,

Davidse 34006 , sessile gland

on pedicel, X 310; B, Pen-

taschistis aspera, Ellis 5439,

sessile gland on leaf margin,

X 310.

copiously, the volatile substance may be an anti-herbivore

mechanism. It is possible that in Prionanthium the glands

produce a sticky substance that would allow mature

spikelets with caryopses to be dispersed through ad-

hesion to animals. I could not investigate this matter

because all populations that I observed in the field were

too young.

Inflorescences in Prionanthium display a distinct re-

duction series. The inflorescence of P. dentatum is the

most unspecialized. It is a condensed panicle that is

superficially spike-like. The short panicle branches are

tightly appressed to the axis of the panicle. In P. ecklonii

most of the inflorescence branches are reduced to pedicels

only and they occur in pairs, one short-pedicelled (near-

ly sessile) the other longer-pedicelled. Only at the base

of vigorous inflorescences is one order of elongated in-

florescence branches developed, and the spikelets are

paired on these branches, except for the apex, as they

are on the main rachis of the inflorescence. Towards the

apex of the inflorescence the paired arrangement is lost

and the spikelets are solitary. In P. pholiuroides the

solitary arrangement of the spikelets is maintained

throughout the inflorescence. In very depauperate speci-

mens of this species the inflorescence may be reduced to

a few, or exceptionally, to a single spikelet.

Except for the non-disarticulating rachis, the structure

of a P. ecklonii inflorescence superficially resembles a

typical andropogonoid raceme, in that the spikelets are

paired and borne unilaterally. As inmost Andropogoneae

the glumes have a much firmer texture than the lemmas

and there are two functional florets. The inflorescence

of P. pholiuroides is clearly even further reduced.

The existence of this transitional series of inflores-

cence types in a small genus is remarkable and clearly

demonstrates how racemose inflorescence types in the

Poaceae can readily arise through a series of progressive

reductions from the paniculate condition. That this has

happened repeatedly in the Poaceae is evident from the

fact that of the 40 tribes recognized in the family by

Clayton & Renvoize (1986), 24 have a racemose type of

inflorescence in at least one of their constituent species.

The lodicules of Prionanthium are fleshy, cuneate,

shallowly one or three-lobed, lack microhairs or cilia

at the apex (Figure 4A— D), and generally have two vas-

cular bundles. They clearly fall within the variation range

encountered in the Arundineae (Tomlinson 1985; Clay-

ton & Renvoize 1986). In P. dentatum (Figure 4 A) the

lodicules have a more prominent lateral lobe than those

in P. pholiuroides (Figure 4B) and P. ecklonii (Figure

FIGURE 4. — Lodicules of Prio-

nanthium species. A, P. den-

tatum, Davidse 33394, in-

ner and outer view of a

pair of lodicules with cut-

off, hairy lemma at the

base, X 150; B, P. pholiu-

roides, Spies 3678, inner

view of old, slightly dis-

torted lodicule, X 150.

C-D, P. ecklonii, Davidse

34018: C, inner view of

lodicule, X 175; D, pair

of lodicules at the base

of the palea with lemma

and ovary cut off, X 85.

Bothalia 18,2 (1988)

147

4C, D), and, in comparison to these two species, their

point of attachment is further away from the palea, i.e.,

they are positioned more toward the dorsal side of the

lemma. The lodicules of the latter two species are very

similar.

The fruit of Prionanthium has never been described.

That of P. dentatum still remains unknown but in both

P. pholiuroides (Figure 5A, B) and P. ecklonii (Figure

5C, D) it is a caryopsis with the pericarp fused to the

seed coat. It is narrowly lanceolate in outline, slightly

convex on the embryo side and grooved on the hilum

side. The hilum is linear. In P. pholiuroides the hilum

groove is nearly as long as the caryopsis and is uniformly

deep (Figure 5B). In P. ecklonii the hilum groove is V2— ■ */„

as long as the caryopsis and is deep in the lower half of

the caryopsis but rather shallow in the upper half (Figure

5D). The embryo is V4-3/10 as long as the caryopsis. The

caryopsis is held loosely between the lemma and palea.

In all respects the caryopsis morphology is typical of

arundinoids.

The manner is which the spikelet disarticulates at

caryopsis maturity is, however, not entirely typical of

arundinoids. As is the norm in the subfamily, disarticu-

lation takes place between the florets, and the glumes

are long-persistent. However, the florets containing the

caryopsis are not shed immediately but each of the two

florets of a spikelet is held between the incurved margins

of the subtending glume. Although the process of dis-

articulation has not been observed beyond this point in

the field, inferences from mature plants of P. ecklonii

and P. pholiuroides which contained a high percentage

of mature caryopses in their spikelets, suggest that the

following events are probable: the florets containing the

caryopses remain enclosed within the glumes for a long

time after the caryopses have matured and after the

plants have died. Presumably the glumes break off

simply due to mechanical wear as the plants disintegrate

during the dry season. Mature inflorescences, with most

spikelets bearing caryopses, which were soaked in water

did not show any noticeable hygroscopic activity of the

glumes or florets under laboratory conditions. Further-

more, it is likely that the upper floret breaks off first

since the upper glume is very prominently hinged and

breaks off more easily than the lower one. This process

seems consistent with the tougher, chartaceous nature

of the glumes compared with the delicate, hyaline florets,

and this suggests that the function of protecting the

caryopsis has been taken over from the lemmas and

paleas by the glumes.

CYTOLOGY

No chromosome counts have ever been reported for

any species of the genus. Young inflorescences were col-

lected in Camoy’s fixative in the field. Anthers were

squashed in aceto-carmine to which a small amount of

iron-acetate was added. Photomicrographs of chromo-

some complements were made with a Reicherdt Univar

microscope.

All species had n = 7 with regular bivalent pairing and

no meiotic irregularities (Table 1; Figure 6A— D). The

only chromosomal irregularity was the occurrence of a

small B-chromosome in one sample ( Davidse 34048 ) of

P. pholiuroides (Figure 6E, F). Besides its small size, it

stained slightly lighter and behaved differently from the

normal bivalents (Figure 6E, F), three characteristics

that define B-chromosomes (Jones & Rees 1982). The

B-chromosome did not line up on the metaphase I plate

(Figure 6F), but since no micronuclei were observed in

the tetrads at the end of the second division it appears

that the B-chromosome is randomly incorporated into

any one of the tetrad nuclei. Due to a paucity of fixed

material in the right stage of meiosis, it was not possible

to study the behaviour of the B-chromosomes at the two

meiotic anr bases nor in pollen meiosis. Although B-

FIGURE 5. — Caryopses of Prio-

nanthium species. A-B,

P. pholiuroides, Ellis 5482A :

A, embryo side; B, hilum

side. C-D, P. ecklonii,

Ellis 5503 A: C, embryo

side; D, hilum side. All

X 17.

148

Bothalia 18,2(1988)

TABLE 1. — Chromosome numbers of Prionanthium species

Species

Chromosome no.

Locality and voucher

P. dentatum (L. f.) Henrard

P. ecklonii (Nees) Stapf

P. pholiuroides Stapf

n=7 CAPE. — 3319 (Calvinia): 16 km S of Nieuwoudtville (-AC), Davidse 33394.

n = 7 CAPE. — 3218 (Clan william): 45 km N of Citrusdal (-BD), Davidse 34018,

Spies 3693.

n=7 CAPE. — 3318 (Cape Town): 17 km W of Malmesbury (-BC), Davidse 33983,

Spies 36 78.

n = 7 + IB CAPE. — 3420 (Bredasdorp): 6 km N of Struisbaai (-CA), Davidse 34048.

chromosomes are generally considered to be genetically

inert, they often affect fertility, especially when present

in high numbers (Jones & Rees 1982). Nevertheless,

their occurrence in the population with the most variable

spikelet morphology is noteworthy. Although this varia-

tion is probably due to simple segregation and recombi-

nation, the possibility of a B-chromosome effect cannot

be entirely ruled out without further research.

FIGURE 6. — Meiotic chromo-

somes of Prionanthium spe-

cies. A-B, P. dentatum,

Davidse 33394, n = 7 : A,

diakinesis; B, metaphase

II. C, P. ecklonii, Davidse

34018, diakinesis, n = 7.

D-F, P. pholiuroides'. D,

Davidse 33983, anaphase I,

n = 7; E-F, Davidse 34048 ;

E, diakinesis, n =7 + IB;

F, metaphase I, B-chromo-

some excluded from the

metaphase plate. Arrows

point to B-chromosome.

Bothalia 18,2 (1988)

149

GENERIC RELATIONSHIPS

Although Chippindall (1955) expressed reservations

about the classification of Prionanthium in the Danthon-

ieae (= Arundineae, cf. Clayton & Renvoize 1986), it has

not been challenged since that time (De Wet 1956; Ren-

voize 1981; Watson et al. 1986; Clayton & Renvoize

1 986). 1 agree with this assessment based on data from leaf

anatomy, gross morphology and chromosome number.

Leaf anatomy has previously been studied by De Wet

(1956) and Renvoize (1981, 1986) in P. pholiuroides

and Watson et al. (1986) in an unspecified species. (Al-

though De Wet (1956) noted that he had studied P. eck-

lonii anatomically, the specimen that he cites as a voucher

apparently is one of the Duthie collections made in the

Stellenbosch Flats and actually represents P. pholiuroides.

His reference to ‘type’ is inexplicable if interpreted as

‘type collection’ since none of the Prionanthium type

collections originated in the Stellenbosch area. Perhaps

he meant to indicate the kind or ‘type of P. ecklonii

originating in Stellenbosch’, which, as I have just indi-

cated, represents P. pholiuroides .) A detailed study of

leaf blade anatomy will be presented by Ellis (in prep.).

From the published results it is clear that Prionanthium

is typically arundinoid in its anatomy. It has anon-Kranz

leaf anatomy that typifies most arundinoid genera, in-

cluding double bundle sheaths, non-radiate mesophyll

with a maximum lateral cell count greater than four,

adaxial ribs, bulliform cell groups not associated with

colourless cells, finger-like microhairs with tapering

distal cells, domed subsidiary cells and sinuous long cells

in the abaxial epidermis, and dumbbell-shaped to nodular

silica bodies. Two unusual features for the subfamily

were noted by Renvoize (1986): 1, the chlorenchyma

extends between the outer bundle sheath and upper

sclerenchyma girders or between upper and lower scle-

renchyma girders, a feature also noted by Watson et

al. (1986); 2, the upper and/or lower epidermal cells

are large and thin-walled. Although unequivocally arun-

dinoid, the anatomical evidence available to date is not

sufficient for a detailed comparison with other possibly

related genera.

All Prionanthium species have a ciliate ligule, a

characteristic that is shared with all other arundinoid

genera recognized by Clayton & Renvoize (1986).

The relationship of Prionanthium to other arundinoid

genera has only been explicitly discussed by Clayton &

Renvoize (1986: 165), who consider it to be one of the

primitive arundinoid genera, along with Tribolium,

Urochlaena, Elytrophorus, Spartochloa, Notochloe, Zen-

keria, Piptophyllum, and Styppeiochloa. They based

this assessment on the short glumes and multinerved

lemmas with entire tips of these genera. Furthermore,

they (Clayton & Renvoize 1986: 171) noted that ‘P.

dentatum hints at a distant relationship with Tribolium ’,

without indicating on which character(s) they made this

judgement. Presumably they suggested this relationship

because P. dentatum has the most fully developed panicle

of the three species and its proportionally short and broad

spikelets superficially resemble those of the Lasiochloa

alliance of Tribolium.

Four spikelet characters point to a relationship be-

tween Prionanthium and Pentaschistis : the occurrence

of well differentiated multicellular glands, two florets

per spikelet, a small rachilla extension above the upper

floret, and similar paleas. Since multicellular glands are

rare in the family, I consider this character especially

important in relating Prionanthium to Pentaschistis.

Although Prionanthium and Pentaschistis share a

number of features, their relationship is not a close one,

as many differences in spikelet characteristics demon-

strate. Of the three characters explicitly mentioned by

Clayton & Renvoize (1986) as typical of the primitive

Arundineae, Prionanthium does not differ strongly from

Pentaschistis in two: 1, short glumes (in both genera the

glumes are as long as the spikelet and are as long as or

longer than the uppermost floret); 2, awnless lemmas

(all species of Prionanthium , 10 out of ±65 species of

Pentaschistis ); 3, multinerved lemmas (3— 5-nerved in

Prionanthium and 5— 11-nerved in Pentaschistis).

The primary basic chromosome number of the Arun-

dinoideae has been considered to be x = 12 (Clayton &

Renvoize 1986). However, it is more likely that this is a

secondary base number derived by polyploidy, since a

number of arundinoid genera are now known with n = 6:

Tribolium (Davidse et al. in preparation, not x = 7 as in-

correctly reported by De Wet 1960 ), Merxmuellera and

Karroochloa (as Danthonia, De Wet 1954, 1960; Du

Plessis & Spies 1988; Spies & Du Plessis 1988), Chinoch-

loa (as Danthonia, Singh & Godward 1963), Schismus

(numerous reports, the latest Faruqi & Quraish 1979; Du

Plessis & Spies 1988 and Spies & Du Plessis 1988) and

Pseudopentameris (Spies & Du Plessis pers. comm.).

Stebbins (1956) and Hunziker & Stebbins (1987) also

consider x = 6 to be the basic chromosome number of

the Arundinoideae. Less common base numbers in the

subfamily are x = 7 and 13 in Pentaschistis (Tateoka

1965; Hedberg & Hedberg 1977). Prionanthium with

n = 7 has a number that is unusual for arundinoids but

one that is matched at the diploid level, n = 7, in part of

the genus Pentaschistis (De Wet 1954, 1960; Du Plessis

& Spies 1988; Spies & Du Plessis 1988). There is also a

resemblance between the two genera in the size of the

chromosomes, among the largest chromosomes in the

subfamily.

CONSERVATION STATUS

All three species of Prionanthium have been included

in the South African red data book (Hall & Veldhuis

1985). The status of both P. ecklonii and P. dentatum

(as P. rigidum ) is listed as uncertain, whereas P. pholiu-

roides is listed as endangered. From the fieldwork and

herbarium research that I conducted, it is apparent that

P. pholiuroides is the most widely distributed of the

three species (Figure 7). The population of P. pholiu-

roides reported on by Hall & Veldhuis (1985) was re-

visited on 21 September, 12 October and 16 November

1987 and still existed as a population of thousands of

individuals. However, the local distribution was very

spotty and clumped. Similar sites only a few kilometers

away lacked any sign of P. pholiuroides. It had also

managed to establish itself in a small area of disturbance

along the roadside. The main population grows in a

heavily over-grazed depression invaded by introduced

species of Acacia.

150

Bothalia 18,2 (1988)

FIGURE 7. — Distribution of Prionanthium dentatum, ★ ; P.

ecklonii, ♦; P. pholiuroides, •.

We discovered a modest range extension for this

species ( Davidse 34048, 34053). Its total distributional

range, although very disjunct, is approximately 250 km,

and not 60 km as noted by Hall & Veldhuis (1985). At

this new site, the species grew on a limestone outcrop,

especially in small sandy soil pockets on the eroded

limestone. This indicates that its habitat requirements

may not be quite as specialized as believed by Hall &

Veldhuis (1985). It seems quite probable that the spe-

cies has been largely eliminated from localities such as

the Stellenbosch Flats, which have been extensively

urbanized. It was common there from the 1920s to at

least the 1940s, judging by the Duthie and Rehm col-

lections, and although I believe it likely that some plants

may still exist in the flats between Gordon’s Bay, Cape

Town and Stellenbosch, these are likely to be so few and

isolated that the species must for practical purposes be

considered extinct in this area. Thus, although the species

is on the whole rare, it may not be in immediate danger

of extinction.

TAXONOMY

Prionanthium Desv., Opuscules sur les sciences

physiques et naturelles: 65, t.4, fig. 3 (1831); Stapf: 455

(1899); Chippindall: 271 (1955); R.A. Dyer 2: 833

(1976); Watson et al.: microfiche 3 (1986); Clayton &

Renvoize: 170 (1986). Type: P. rigidum Desv. =P. den-

tatum (L. f.) Henrard.

Prionachne Nees in Lindley: 447 (1836). Chondro-

laena Nees: 133 (1841), nom. superfl. pro Prionachne.

Type: P. ecklonii Nees = Prionanthium ecklonii (Nees)

Stapf.

Annuals. Leaves linear; ligule a ciliate membrane. In-

florescence a spike-like panicle or a secund raceme or spike

bearing solitary or paired, subsessile or short-pedicellate

spikelets. Spikelets 2-flowered with a small rachilla ex-

tension, laterally compressed, keeled; disarticulation

above the glumes and between the florets. Glumes sub-

equal, with the lower slightly longer than the upper, as

long as the spikelets, slightly asymmetrical, keeled,

rigidly chartaceous and 5— 8-nerved in the centre, mem-

branous to hyaline marginally, long persistent after

spikelet maturity and each individually enclosing one of

the florets at caryopsis maturity; bases coriaceous and

hinged, the upper more prominently hinged than the

lower; keels usually with prominent, multicellular glands.

Florets bisexual. Lemmas hyaline, 3— 5-nerved, subequal,

nearly as long as or slightly shorter than the glumes,

covering the palea wings but not the palea keels at spike-

let maturity. Paleas nearly as long as or slightly shorter

than the lemmas, 2-nerved and 2-keeled, the back and

wings hyaline, narrow, the keels prominent, thickened,

well developed, exposed at spikelet maturity. Lodicules

2, cuneate, fleshy, glabrous, shallowly 1- or 3-lobed.

Stamens 3. Ovary glabrous; styles 2, separate; stigmas

plumose, laterally exserted. Caryopsis narrowly lanceo-

late in outline, convex on the embryo side, shallowly

grooved on the hilum side, especially toward the base;

hilum linear; embryo ± '/4 as long as the caryopsis.

By contrast, P. dentatum and P. ecklonii appear to

have a much more limited distribution and on this basis

must be considered to be endangered.

Not surprisingly for annual species, samples of P. pho-

liuroides {Ellis 5482 A) and P. ecklonii {Ellis 5503A)

collected in November 1987 showed good caryopsis

production. In both species, at random, five spikelets

from five different plants were examined for caryopsis

production in both the upper and lower florets. In P.

pholiuroides the lower florets had 92% caryopsis pro-

duction and the upper 88% . In P. ecklonii 96% of the

lower florets and 92% of the upper florets produced

caryopses.

All three species appear to be able to exist in areas of

moderate disturbance such as that encountered in the

typical grazed veld of the Cape. All three species also

grew on roadsides which may actually undergo less dis-

turbance than adjacent grazed veld since animals general-

ly do not have access to roadsides. However, road con-

struction projects could have drastic effects on these

small roadside populations since the vegetation may be

completely scraped off by heavy machinery at any time.

A genus of three species endemic to the Cape Province.

Key to species

la Florets pubescent 1 . P. dentatum

lb Florets glabrous:

2a Spikelets solitary; inflorescence usually unbranched;

glands on glume keels sessile or slightly stalked,

rarely absent 3. P. pholiuroides

2b Spikelets paired except solitary near the apex of the

inflorescence; inflorescence (except in depaupe-

rate specimens) with appressed branches at the

base; glands on glume keels conspicuously stalk-

ed 2. P. ecklonii

1. Prionanthium dentatum (L. f) Henrard in

Blumea 4: 530 (1941). Type: Cape, Bockland, 1773,

Thunberg s.n. (UPS, holo.— PRE, microf.!; PRE, frag-

ment!; BM!,K!,MO!).

Phalaris dentata L. f.: 106 (1781); Thunberg: 19 (1794).

Phleum dentatum (L. f.) Pers. 1: 79 (1805). Chilochloa dentata

(L. f.) Trin.: 168 (1824), nom. nud., 1: t. 73 (1827). Prionan-

thium rigidum Desv.: 65 (1831); Stapf: 455 (1899). Lasiochloa

pectinata Trin.: corr. & emend, t. 73 (1836), nom. illeg. superfl.

pro Phalaris dentata L. f. Chondrolaena dentata (L. f.) Steud. :

355 (1840). Prionachne dentata (L. f.) Nees: 134 (1841), pro

syn. Chondrolaena phalaroides Nees.

Bothalia 18,2 (1988)

151

Culms 30-430 mm tall, unbranched or branched

only at the base, glabrous. Sheaths short-pubescent;

ligule 0,5- 1,4 mm long, the membrane 0,05—0,2 mm

long, the cilia 0,4— 1,3 mm long; blades 15-105 x 0,5—

3,0 mm, short-pubescent. Inflorescence a condensed,

cylindrical spike-like panicle, 5-75 mm long, reduced to

a single spikelet in depauperate individuals; rachis cylin-

drical, pubescent; branches tightly appressed to the

rachis, shortened upward; pedicels short stumps or to 1

mm long. Spikelets 3,2— 5,2 mm long, not obviously

borne in pairs, laterally arranged; glumes 5— 7 -nerved,

the side facing the rachis pubescent (Figures 1 A, 2C), the

hyaline margin broader than the chartaceous centre, the

keel with prominent stalked glands, the tip shortly awn-

pointed; lemmas evenly pubescent, 3-nerved, shortly

awn-pointed; paleas glabrous on the margins, pubescent

on the back; anthers 1,8— 2,3 mm long. Chromosome

number : 2n = 14 (Figure 6A, B).

Only known from the Nieuwoudtville area of the

western Cape at an elevation of ± 700 m in Western

Mountain Karoo (Figure 7).

Vouchers: Davidse 33394, 33396; Ellis 2452, 5416, 5417.

Nees (1841) and Chase & Niles (1962), among others,

considered Thunberg’s (1794) Phalaris dentata to be new

and different from the Phalaris dentata of Linnaeus

(1781). This is incorrect because both names are based

on Thunberg’s collection from ‘Bockland’ (= Bokkeveld).

This is clear from Juel’s (1918) account explaining how

Linnaeus published hundreds of new species based on

Thunberg’s Cape collections. I interpret Thunberg’s usage

to be based on Linnaeus’s (1781) name without proper

attribution and using a different and original descrip-

tion. For this reason, I consider all other combinations

made with the epithet dentatum to be based on Lin-

naeus’s (1781) initial usage of this name.

Until Ellis re-collected it in 1975, this species was

only known from the type collection made by Thunberg

in 1773. Thunberg’s collection, however, was a large one

and a number of duplicate specimens were distributed

to other herbaria so that very good type material exists.

2. Prionanthium ecklonii (Nees) Stapf in Flora

capensis 7: 456 (1899); Chippindall: 271 (1955). Type:

‘C. b. sp.’ (Cape of Good Hope). Lectotype (chosen here):

ad Olifantsrivier fluviam alt. I, Clan william, Ecklon s.n.

(MO!; BM, isolecto.!; PRE, fragment!; US, isolecto.!).

Prionachne ecklonii Nees in Lindley: 448 (1836). Chondro-

laena phalaroides Nees: 134 (1841), nora. illeg. superfl. pro Pri-

onachne ecklonii. Chondrolaena phalaroides Nees var. dentata

Nees: 134 (1841).

Culms 190—370 mm tall, usually branched, purple,

glabrous. Sheaths puberulous or glabrous, usually ciliate.

Ligule 0,3— 0,6 mm long, the membrane 0,05-0,1 mm

long, the cilia 0,3— 0,5 mm long; blades 40-160 x 0,5—

1,5 mm, glabrous or appressed pubescent. Inflorescence

an inconspicuously secund spike-like raceme or panicle,

15—95 mm long; rachis cylindrical, glabrous to sparsely

puberulent; branches 30 mm long, tightly appressed to

the main rachis; pedicels unequal stumps. Spikelets

4;4_6,1 mm long, usually borne in pairs in the middle

part of the inflorescence, solitary toward the apex of

the inflorescence and its branches, laterally arranged,

the pairs borne alternately; glumes 5— 8-nerved, the side

facing the rachis pubescent, narrower, 1— 3-nerved, the

side away from the rachis usually broader and glabrous,

sometimes sparsely pubescent, 3— 4-nerved, the midrib

with prominent, stalked multicellular glands; lemmas

hyaline but herbaceous at the tip, sometimes scaberu-

lous on the midrib, otherwise glabrous, 3-nerved; paleas

glabrous, usually bilobed at the apex, sometimes sparsely

ciliate at the apex; anthers 2,9— 3,9 mm long; caryopsis

2,5— 3,2 mm long, the hilum V2 — 4/s as long as the cary-

opsis (Figure 5D). Chromosome number'. 2n = 14 (Figure

6C).

Only known from the south-western Cape Province at

low elevations, rare, in Coastal Renosterveld (Figure 7).

Vouchers: Davidse 34018; Drege s.n.; Ecklon s.n.; Ecklon

& Zeyher s. n. ; Spies 36 93.

This species was until recently only known from three

collections, two made by Ecklon and one by Drege.

When Nees (in Lindley 1836) described Prionachne eck-

lonii, he did not indicate a type and only a general geo-

graphical distribution was provided. However, from his

use of Ecklon’s name as the epithet and the fact that all

three of the collections had been available to him by

1836 (Gunn & Codd 1981), it is clear that he was basing

his species on these collections. This is confirmed from a

later publication (Nees 1841), dedicated to Drege, Ecklon

and Zeyher, where he redescribed and renamed this

species as Chondrolaena phalaroides with two varieties,

var. dentata and var. edentula, and explicitly cited the

Ecklon and Drege collections. He also referred to his

1836 treatment of Prionachne in the synonymy. Unfor-

tunately he did not directly refer to P. ecklonii but made

the new, but illegitimate, combination Prionachne den-

tata. This new synonym was based on Linnaeus’s (1781)

Phalaris dentata, which was also cited in synonymy, but

was in fact misapplied. The fact that Nees explicitly

states that he was publishing a substitute name for his

Prionachne indicates that he considered Chondrolaena

phalaroides and Prionachne ecklonii to be the same

species. Accordingly I have chosen to lectotypify Prio-

nanthium ecklonii with the Ecklon collection from the

Olifantsrivier. This preserves the usage that Stapf (1899)

initiated in his excellent treatment of the genus in Flora

capensis.

Nees (1841) did not truly understand the differences

between the two species that had been collected by this

time. This is apparent from the fact that he synonymiz-

ed Phalaris dentata (= Prionanthium dentatum ) with his

new species Chondrolaena phalaroides (= Prionanthium

ecklonii ). When he renamed Prionachne ecklonii as

Chondrolaena phalaroides, he recognized two varieties,

var. dentata and var. edentida. Under var. dentata he

listed all but one of the combinations and usages of this

name, including his own illegitimate new combination,

Prionachne dentata. He excluded Thunberg’s (1794,

1813) interpretation of Phalaris dentata because he felt

that Thunberg’s description, which was substantially

■different from that of Linnaeus (1781), could not apply

to the same species. In fact, Thunberg’s descriptive state-

ments queried by Nees, spike ‘subpaniculatum’ and

'glumas ovatas’, do apply to Prionanthium dentatum,

especially in contrast to the inflorescence and glume

152

Bothalia 18,2 (1988)

shapes of P. ecklonii. Actually, Thunberg’s entire glume

description (glumae ovatae, concavae, glabrae, bimargi-

natae vel cinctae, intra marginem linea duplici aut tri-

plici elevata, viridi, sinuato-serratae) applies very well

to the two glumes considered as a single unit. Further-

more, Thunberg (1794) explicitly cites the locality of

the collection (crescit in Bockland) which leaves no

doubt that it was based on his own collection from the

Cape, which was, as noted before, also studied by Lin-

naeus (1781).

Nees’s var. edentula should be referred to Prionan-

thium pholiuroides (see the discussion following that

species).

3. Prionanthium pholiuroides Stapf in Flora

capensis 7: 456 (1899). Chippindall: 271 (1955). Type:

Cape, Fish Hoek Valley, damp hollow, Nov. 1897,

Wolley Dod 3394 (K, holo.!; BM!, BOL!, MO!, PRE!).

Chondrolaena phalaroides Nees var. edentula Nees: 134

(1841). Type: not located, probably destroyed at B.

Prionachne pholiuroides (Stapf) Phillips: 6, t.63 (1931).

Culms 40—250 mm tall, usually branched, purple,

glabrous. Sheaths glabrous; ligules 0,3— 0,9 mm long,

the membrane 0,1— 0,3 mm long, the cilia 0,2— 0,7 mm

long; blades 15—70 x 0,5— 1,5 mm, glabrous below;

scaberulous above. Inflorescence a secund, 2-ranked

spike, 15—60 mm long; rachis triquetrous, curved out-

ward at spikelet insertion, glabrous to scaberulous; pedi-

cel reduced to a stump. Spikelets 3,1— 7,0 mm long, the

terminal one longest, usually borne singly and alternate-

ly on the rachis, the lateral side of the lower glume of

each spikelet appressed to the rachis; glumes 7— 8-nerved,

the midrib and sometimes some or all the nerves with

prominent sessile glands, rarely eglandular with prickles

on the keels (Figure 1G), glabrous, except pubescent on

the inner side of the lower glume, rarely also puberulent

on the outer side of the lower glume or completely

glabrous; lemmas hyaline but herbaceous in the middle

toward the apex and slightly scabrous, otherwise glabrous,

the lower 3-5-nerved, the upper 3-nerved; paleas gla-

brous; anthers 1,8-3, 5 mm long. Caryopsis 2, 5-3,0

mm long, the hilum nearly as long as the caryopsis

(Figure 5B). Chromosome number : 2n = 14 and 14 + IB

(Figure 6D— F).

Only known from the Cape Province, at low eleva-

tions, rare, primarily in shallow depressions that tempo-

rarily collect water (Figure 7).

Vouchers: Anderson 8; Davidse 33983; Duthie 1767; Ellis

5434; Rehm 6054.

When Nees (1841) described Chondrolaena phalaroi-

des var. edentula, he described the glumes as non-dentate

(i.e., lacking stalked, multicellular glands) and the spike-

lets as entirely alternate (i.e., spikelets borne singly on

the rachis). This description applies well to Prionanthium

pholiuroides, especially when contrasted to P. ecklonii.

Nees also noted that he only saw a fragment of this

taxon, intermixed with other specimens. Presumably

this fragment was included with one of the three cited

specimens, although he does not specify which one. All

three were presumably destroyed since all his Poaceae

specimens were acquired by B in 1855 (Stafleu & Cowan

1981). I have not seen similar fragments or specimens

among duplicates of the Ecklon and Drege collections

that I have examined. Nees suspected it to represent a

new species, but, presumably because of the fragmentary

nature of the specimen, decided to recognize it at a

lower rank.

A recently discovered population near Struisbaai

( Davidse 34048, 34053 ) is quite variable in spikelet

morphology. It has the smallest spikelets of all known

populations, 3, 1-5, 2 mm versus 4, 5-7,0 mm for all

others. However, more importantly, individual plants

within the population have spikelets with glumes that

vary from completely eglandular and glabrous to those

that are typically glandular and pubescent on the inner

side, to glandular-glabrous, or eglandular-pubescent.

Since these plants are seemingly segregating for these

characters, these variations are not taxonomically re-

cognized. The Malmesbury population represented by

Davidse 33459 is variable with regard to glabrous and

pubescent spikelets. The only population, apparently

now extinct, with glands on nerves other than the mid-

nerve of the glumes, is the Stellenbosch Flats population

sampled by Duthie from 1925 to 1928.

ACKNOWLEDGEMENTS

I thank the Department of Agriculture and Water

Supply for the fellowship which enabled me to conduct

this research in South Africa and the Botanical Research

Institute, for research facilities. I am grateful to B. de

Winter and M. de Winter for their very generous hospital-

ity during my time in South Africa. I thank R.P. Ellis

and J.J. Spies for their helpful assistance in all aspects of

my stay in South Africa. I also thank S.S. Brink, G.

Condy, J. Davidse, H.duPlessis, J.D.Dwyer, H.Ebertsohn,

A. Fellingham, L. Fish, R.E. Gereau, G.E. Gibbs Russell,

H.P. Linder, E.G.H. Oliver, S.M. Perold, C. Reid for

assistance in the fieldwork, research, and preparation of

the manuscript. I thank the curators of BM, BOL, K,

MO, PRE, STE, and US for making their specimens avail-

able for study.

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23: 1-97.

CHASE, A. & NILES, C.D. 1962. Index to grass species. Vol. 3.

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1: 78-136.

SPECIMENS EXAMINED

Adamson 3268 (3) BOL, BM, NBG; Anderson 8 (3) BOL, K,

MO, PRE, STE.

Davidse 33394 (1) BOL, MO, NBG, PRE; 33396 (1) MO, PRE;

33459 (3) BOL, MO, NBG, PRE; 33983 (3) BOL, MO, NBG,

PRE; 34018 (2) BOL, MO, NBG, PRE; 34048 (3) BOL, MO,

NBG, PRE; 34053 (3) BOL, MO, NBG, PRE; Drege s.n. (2)

BM, K, MO, NBG, PRE; Duthie 176 7 (3) BOL, PRE, STE; 1 795

(3) PRE, STE; s.n. (3) PRE.

Ecklon s.n. (2) BM, MO, NBG, PRE, US; Ecklon & Zeyher s.n.

(2) PRE; Ellis 2452 (1) PRE; 2453 (1) K, PRE; 5416 (1) PRE;

5417 (1) PRE; 5433 (3) PRE; 5434 (3) PRE; 5435 (3) PRE;

5482A (3) MO; 5503A (2) MO; Esterhuysen s.n. (3) BOL.

Parker 3745 (3) BOL, K, NBG.

Rehm 6054 (3) BM, K, PRE; Ronaasen s.n. (3) NBG.

Spies 36 78 (3) PRE; 3693 (2) PRE.

Thunberg s.n. (1) BM, K, MO, PRE.

Wolley Dod 3394 (3) BM, BOL, K, MO, PRE.

Zeyher s.n. (as Pappe s.n.) (3) BOL, NBG, PRE.

Note added in proof

Stalked, multicellular glands also occur on the back

of the lower lemma in many species of Panicum sect.

Stolonifera [Zuloaga, F.O. & Sendulsky, T. 1988. A revi-

sion of Panicum subgenus Phanopyrum section Stoloni-

fera (Poaceae: Paniceae). Annals of the Missouri Botani-

cal Garden 75: 420— 455] of tropical America. Although

the walls of the stalk are similar to those of Prionanthium,

the central depression is nearly smooth and lacks the

rectangular grid. It is probable that these Panicum glands

are secretory, but this has not yet been conclusively

demonstrated.

Bothalia 18,2: 155-163 (1988)

Studies in the genus Riccia (Marchantiales) from southern Africa. 10.

Two new white-scaled species of the group ‘Squamatae’: R.

argenteolimbata and R. albornata

O.H. VOLK*, S.M. PEROLD** and T. BORNEFELD***

Keywords: Marchantiales, new species, nothopolyploidy, Riccia, southern Africa, taxonomy

ABSTRACT

Two new white-scaled species, R. argenteolimbata and R. albornata have been isolated by one of us (Volk).

R. argenteolimbata has a compact thallus with stiff, regular scales and apolar spores, whereas R. albornata has a

somewhat more spongy structure of the thallus, large, frilly, hyaline scales and polar spores.

UITTREKSEL

Twee nuwe spesies met wit skubbe, R. argenteolimbata en R. albornata is deur een van ons (Volk), gei'soleer.

R. argenteolimbata het ’n kompakte tallus, met stywe, ewe-groot skubbe en apolere spore, terwyl R. albornata

’n ietwat meer sponserige tallus-bou, groot, gekartelde, hialiene skubbe en polere spore het.

1. Riccia argenteolimbata Volk & Per old, sp. nov.

Thallus dioicus, perennis, parvus, in vivo glaucescens-

opacus, in sicco albidus; squamis argenteis quasi limbatis,

inde nomen. From usque ad 7 mm longa, 0,7— 1 ,2(— 2,0)

mm lata, 0,6— 0,9 mm crassa, 1 — 1,5(— 2)-plo latior quam

crassa, asymmetrice furcata, lobis extremis brevibus,

obovato-ligulatis, subacutis, marginibus acutis, lateribus

costae rectis; pagina superior subplana, longe sulcata.

Squamae rigidae, dense imbricatae, semiorbiculatae, inte-

gerrimae, marginem frondis minutim superantes, calce

incrustatae, albae vel plusminusve subfuscae. Sporae

apolares, subglobosae, sine ala, 80-120(-130) pm dia-

metro, rufo-bruneae, ornamentatione reticulata, 12—15

(-16) foveolis in diametro. Chromosomatum numerus

n = 8; 9; 16; 20; 24.

TYPES.— SWA/Namibia, 2116 (Okahandja): Marien-

hof (Dunroamin) (OK 289) (— BD), alt. 1 550 m, schluff-

reicher, stark kalkhaltiger, grauer Boden der ‘Vlakte’,

zeitweise durchfeuchtet, pH 7,8, 1974.03.28, Volk

00910 (M, holo.). 2217 (Windhoek): Hatsamas (WIN

92) (-DC), alt. 1 870 m, ebene Treppenstufen im Blau-

kalk, staubiger, schluffreicher, kalkhaltiger Boden, pH

7,8, mit Oropetium capense, 1974.01.31, Volk 00 762

(M, para.).

Thallus dioicous (Figure 1 A, B), perennial, gregarious

or single and scattered, rarely in rosettes; mostly asym-

metrically bi- or trifurcate, branches medium to widely

divergent, obovate-ligulate (Figures 1A, B; 2 A), 2—7

mm long, 0,7-l,2(-2,0) mm broad, 0,6-0, 9 mm thick,

1 — 1 ,5( — 2) times broader than thick, segments short,

apex wedge-shaped (Figure 2B); sulcus conspicuous and

long (Figure 1A, B, E), its sides convex, becoming flatter

proximally (Figure 1H1-3); dorsally dull greenish grey,

* Botanische Anstalten d. Univ., Wurzburg D8700, Germany, BRD.

** Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001, RSA.

*** Am Reelein 1, Hochberg D8706, Germany, BRD.

MS. received: 1988.02.01.

not shiny, sometimes brown along the margin; margins

acute, flanks steep, dark-coloured, covered by silvery

grey scales, hence the specific epithet; ventral surface

rounded, green or reddish brown, apically with arched,

narrow, brown bands across (Figure 1G); when dry,

dorsally greyish white, margins lip-like tightly inflexed

and flanks densely covered by regularly arranged, ap-

pressed, stiff scales (Figure 1C). Anatomy of thallus :

dorsal epithelium bistratose, upper layer of cells rarely

intact, but when intact, inflated, ± 20-35 x 30-40 pm,

capped with deposits of calcium, (Figure II), soon col-

lapsed, forming so-called ‘ring’ -cells, which appear thick-

walled (Figures 1 J, centre; 2C, D), the lateral walls

double and raised; second layer of epithelial cells with-

out chloroplasts, short-rectangular, 25-37 x 22—32 gm,

upper transverse and lateral walls thicker, the latter

gradually thinning out below (Figure II); from above,

dorsal cells arranged in regular, honeycomb pattern,

air pores mostly triangular and quadrangular, small,

(Figures 1 J, 2D); assimilation tissue (chlorenchyma)

compact, about '/2 the thickness of thallus, cells rect-

angular, 40—50 x ± 32 pm, in columns of about 8—10

cells, enclosing very narrow, usually 4-sided air canals;

storage tissue almost '/2 the thickness of thallus. Rhi-

zoids 12—23 pm wide, mostly smooth. Scales generally

only marginally, sometimes entirely white, base silvery

mauve becoming dark grey-brown more proximally

(Figure IB, C, E, F, K), closely imbricate, stiff, 600—

800 x ± 500 pm, projecting about 100 pm above thallus

margin (Figure 2E, F), rounded, edge smooth, cell

surfaces heavily encrusted with calcareous deposits

(Figure 1L); in body of scale, cells 50-90 x 25—30 gm,

hexagonal or nearly isodiametric, corners angled, cell

walls appear thick; at margins cells smaller, quadrate to

brick-shaped, ± 22 x 27 pm. Antheridia with short, hya-

line necks. Archegonia with dark purple necks. Sporangia

usually single, bulging slightly dorsally, each containing

200-450 spores, rarely sporulating. Spores 80-120

(-130) gm in diameter, reddish brown to almost black,

semi-opaque to opaque; globular to subglobular, apolar,

wing and triradiate mark absent (Figure 3A, B), peri-

phery tuberculate (Figure 3D, E), ornamentation reticu-

156

Bothalia 18,2 (1988)

FIGURE 1. — Riccia argenteolimbata. Morphology and anatomy. A, female thallus with two sporangia, turgid; B, male thallus

with antheridial necks, turgid; C, as B, dry, scales with dark bases; D, bulbil; E, turgid, scales white; F, same specimen as

E, dry; G, transverse bands on ventral surface; Hl-3, transverse sections from apex to base; I, part of section with de-

posits on turgid and collapsed epithelial cells, (‘ring’ cells with thickened walls); J, horizontal section at different levels:

left, top of epithelium with triangular air pores; centre, base of epithelial cells; right, chlorenchyma with four-sided air

canals; K, scale; L, calcium deposits on outer surface of scale; (A-L by Volk). A, E, G, Volk 00910 ; B, D, H3, Volk

00765\ HI, Volk 84-713; H2, 1-L, Volk 881a. Scale bars: A-H=lmm; I, J, L =50 urn; K = 100 jim.

Bothalia 18,2 (1988)

157

feld & Grillenberger 1987). The same degree of notho-

polyploidy, represented by two different chromosome

patterns within one species, occurs in R. argenteolim-

bata (Figure 4B, C) and in R. albolimbata (Bornefeld

unpublished). The karyotype with n = 20 (Figure 4D)

seems to have developed from the one with n = 24

(Figure 4C) by the loss of the four D, chromosomes.

The loss of a group of homologous chromosomes has

also been observed in Targionia lorbeeriam (Bornefeld

1987). Further investigations may yield still other

karyotypes of R. argenteolimbata . Because of the vari-

ability of chromosome numbers in some Riccia species,

this character does not seem to be suitable for the

definition of a species.

Chromosome numbers (Number of counts in brackets)

n = 8 : Volk 86-930 pp (3), Volk 86-934 (5), S.M.

Perold 772 (3)

n = 9 : Volk 86-930 pp (4), Volk 86-934 (3)

n = 16: S.M. Perold 727 (2)

n = 20: Volk 81-164/165 (4)

n = 24: with one A chromosome: Volk 81-164/ 165

(8), Volk 81-1 70 (2), Volk 81-1 72 (3), Volk

86-922 (2); with two A chromosomes: Volk

81-204 pp (2); Volk 00912 (5)

FIGURE 2. — Riccia argenteolim-

bata. Morphology and ana-

tomy. A, dorsal view of

dichotomous branch; B,

wedge-shaped apex and

groove; C, dorsal cells with

only a few not collapsed;

D, dorsal cells collapsed,

showing cell walls and small

pores; E, stiff scales at

thallus margin; F, more

enlarged view of marginal

dorsal cells and scales. (SEM

micrographs by Perold).

(A-F, S.M. Perold 772).

Scale bars on A-F =100

/urn.

late with 1 2 — 1 5( — 1 6) round to angular areolae across

diameter (Figure 3C, D), 3-7 Atm wide, ridges thick

(Figure 3A, F), raised at nodes into conical or truncate

processes (Figure 3E). Chromosome numbers n = 8; 9;

16; 20; 24 (Figure 4A, B, C, D).

Riccia argenteolimbata is one of eight southern Afri-

can Riccia species for which several karyotypes are

known. Four of the six karyotypes of R. argenteolim-

bata (Figure 4A-D) are given in their original arrange-

ment and sorted according to chromosome types (Borne-

feld 1984). The karyotype with n = 8 (Figure 4A) re-

presents the basic set in the genus Riccia and is common

to the majority of the species. The karyotype with n = 9

possesses one additional B> chromosome; that with n =

16 has two B, , two Cj and four Dj chromosomes. As is

evident from Figure 4B and 4C, the triploid karyotypes

of R. argenteolimbata are not achieved by threefold

multiplication of the basic set, but by differential multi-

plication of the diverse chromosome types. For this

phenomenon, which is unique to Riccia, the term ‘notho-

polyploidy’ was earlier suggested (Bornefeld 1984). The

decreasing size of homologous chromosomes does not

occur by chance; it is also found in the other genera of

the Marchantiales and becomes more noticeable as the

number of homologous chromosomes increases (Borne-

158

Riccia argenteolimbata Volk & Perold, sp. nov., dif-

fers from the other Riccia species with white scales by

being dioicous, by the compact texture of the thallus,

the closely imbricate, stiff, white or greyish white scales,

often with darker coloured bases, and by the globular

or subglobular, apolar spores.

Among the Volk specimens 881, 883 and 11906,

which Arnell (1957) identified as R. albosquamata, thalli

of R. argenteolimbata are also present; however, his de-

scription of R. albosquamata, as well as the type speci-

men Volk 452, and most definitely its triangular-globular,

polar spores, do not correspond with/?, argenteolimbata

(Perold in press). Arnell appears to have been uncertain

about the distinguishing characters of the white-scaled

species and misidentifled them several times.

R. argenteolimbata is found at altitudes of 1 000—

2 000 m above sea level, in summer rainfall areas with an

annual precipitation of at least 200 mm. It prefers a

sunny position, is calciphilous and often grows on grey-

ish turf soils with a high dust content, overlying lime-

rich beds, calcrete, crystalline limestone, or dolomite, or

on alluvium on steep river banks. These soils may be-

come periodically waterlogged for short intervals when it

rains. Soil pH values are high: of 96 samples tested, 36%

fell between 7,0 and 7,4; 39% between 7,5 and 7,9 and

Bothalia 18,2 (1988)

25% above 7,9 (Volk). The species is often associated

with R. trichocarpa, R. albolimbata, R. atropurpurea,

Cyanophyceae and with crustaceous lichens. It has so

far been found only in Zambezian areas occupied by the

Sudano-Zambezian floral element (Volk 1966), in the

northern parts of SWA/Namibia and a few localities in

Botswana, north-western and south-western Transvaal,

central Orange Free State and northern Cape Province

(Figure 5). In some recent collections, e.g. Volk 85-775

and 86-930, very small plants are mixed with plants of

normal size. The difference in size seems to be genetical-

ly determined, as it persists in cultivation experiments

and the small specimens may represent a different sub-

species.

During the dry season from April onwards, R. argenteo-

limbata tends to form bulbils or turions for perennation

and dispersal (Figure ID) (Volk 1984); these are very

loosely attached to the soil, brown material accumulates

in the walls of the peripheral cells, which then form a

resistant covering, the assimilation columns shorten and

fatty oils appear in the storage tissue.

SPECIMENS EXAMINED

S.W.A./NAMIBIA. — 1815 (Okahakana): Etosha, Bitter-

water (-CC), Volk. 01389 (M). 1816 (Namutoni): Olifants-

bad (-AA), Volk 81-174 (M, PRE); Etosha, Okerfontein

FIGURE 3. — Riccia argenteolim-

bata. Spores. A, ?proximal

face; B, side view; C, ?dis-

tal face; D, ?distal face,

with crowding of truncate

papillae at areolar nodes in

centre; E, more enlarged

view of areolae; F, ?distal

face. [A-E, SEM micro-

graphs; F, LM (light micro-

scope) micrograph by Pe-

rold.] (A, B, D, Volk

8 5- 7 73b-, C, E, F, Volk

84-713.) Scale bars on A-E

= 50 Mm; diameter of spore

on F, ± 100 Mm.

Bothalia 18,2 (1988)

159

u

cc

Cc

FIGURE 4. — Riccia argenteolim-

bata. Karyotypes. A, karyo-

type with n = 8 and chro-

mosome formula where A,

BB, CC, DD, E represent

the basic set in the genus

Riccia', B, nothotriploid

set with one A chromo-

some and U2 chromosome

(this karyotype seems to

be the most widespread

one) ex Bomefeld 1984

(by courtesy of J. Cramer

in der Gebriider Bomtrae-

ger Verlagsbuchhandlung,

Berlin-Stuttgart, FRG); C,

nothotriploid set with two

A chromosomes and lack-

ing U2 chromosome; D,

karyotype with n = 20 re-

sembles that of 2C, but the

four Dj chromosomes have

been lost (the sequence

from A to D possibly cor-

responds to an evolution-

ary line). (A-D by Borne-

feld) (A, S.M. Perold 772;

B, Volk 81-J72 p.p.; C,

Volk 81-204 p.p.; D, Volk

81-164/165.) Scale bar:

A— D = 1 /rm.

)C«

ci C2 c'2

« << «

D1 Dg

©<» jr

• i

«

O

c

D

«

I

(-DD), Volk 81-164 (M, PRE). 1915 (Okaukuejo): Etosha

between Okaukuejo and Okandehoe (-AA), E. Retief 1422a

(PRE); Etosha, Sprokieswoud (-BA), Volk 81-172 (M); Okau-

kuejo (-BB), Volk 00912, 81-204 (M). 1916 (Gobaub): Etosha,

Gemsbokvlakte (-AA), E. Retief 1493a (PRE); Smook 5118

(PRE); OU 449 (-AC), Volk 5116 (M); Halali (-BA), Smook

5138 (PRE); Elandsdraai (-CA), Volk 81-170 (M); GR 412

(-CB), Volk 00749 (M); OU 179 (-CD), Volk 00975 (M);

GR 398 (-DB), Volk 00744 (M); OU 359 (-DC), Volk 81-1 77a

(M). 1917 (Tsumeb): Otjikoto Lake (-AB), Smook 5159 (PRE);

Volk 81-156 (M, PRE); GR 45 (-DB), Volk 81-151b (M, PRE);

GR 154 (-DC), Volk 00454 (M). 1918 (Grootfontein): GR 729

(—AC), Volk 81-146 (M, PRE); Volk 84-705 (M). 2014 (Wel-

witschia): OU 419 (-BA), Volk 5137 (M). 2016 (Otjiwarongo):

OU 193 (-AA), Volk 00977 (M); OU 132 (-AA), Volk 5135

(M); Outjo Dist., 20 km W of Khorixas (- AA), Long & Rae 921

(E); OU 187 (—BA), Volk 86-860 p.p. (M); OTJ 23 (-CA),

Volk 00462 (M); OTJ 97 (-CC), Volk 84-713 (M). 2017 (Water-

berg): GR 98 (-AA), Hoffmann PRE-CH 4514 (PRE); Volk

00453, 84-692, 84-721, 85-773, 85-775, 86-930, 86-933, 86-934

(M); OTJ 147 (-CA), Volk 881 p.p., 883 p.p. (M, PRE). 2116

(Okahandja): OM 46 (-AA), Gibbs Russell & Smook 5240

(PRE); OM 37 (-AA), Volk 85-853 (M); OK 240 (-BD), Volk

00910 (M); OK 34/44 (-CA), Volk 11906 p.p. (M); OK 22

(-DA), Volk 5170 (M); OK 252 (-DA), Volk 00469 (M). 2117

(Otjosondu): OK 105 (-DB), Volk 6169 (M). 2118 (Steinhau-

sen): OK 107 (-DB), Volk 86-922 (M); OK 113 (-DB), Volk

6212a (M); GO 252 (-DB), Volk 00507 (M). 2217 (Windhoek):

Avis Dam (-CA), Volk 00752 (M); Rietfontein (-CD), \olk

81-265 p.p. (PRE); WIN 92 (-DC), Volk 00591, 00761, 00762,

00764, 00765 (M). 2316 (Nauchas): Windhoek Dist., Gamsberg

Mountains, 130 lan SW of Windhoek (-AD), Long & Rae 946

(E); WIN 46 (-BA), Volk 81-200 (M). 2317 (Rehoboth): GIB

113 (— BB), Volk 01291 (M). 2416 (Maltahohe): MAL 19

(-BB), Volk 6386, 6439, 6860 (M); MAL 91 (-BD), Volk 6856

(M); 2417 (Mariental): GIB 6 (-BA), Volk 5268 (M). 2418

(Stampriet): Gochas, Auob River (-DD), Hardy 6586a (PRE).

BOTSWANA.— 1921 (Aha Hills): Ngamiland Dist., Aha

Hills, hill on N side near E end of range' (-CC), Long & Rae 834,

160

842 (E). 2123 (Pink Pan); Central Kalahari Game Res., Decep-

tion Pan (-BD), Henderson 659 (PRE).

TRANSVAAL. — 2228 (Maasstroom): Alldays, 55 km W of,

on road to Swartwater (—DA), S.M. Perold 766, 767, 769, 772

(PRE); Gregory Halt, 19 km NW of Alldays, near bridge (-DB),

S.M. Perold 737 p.p. (PRE); opposite Bulkop Store, on calcrete

(-DC), S.M. Perold 793 p.p., 794 (PRE). 2327 (Ellisras): Farm

Franschhoek, 29 NW of Villa Nova (-BD), Smook 4231 (PRE).

2329 (Petersburg): Vivo, 15 km N of (-AB), S.M. Perold 727,

728 (PRE). 2428 (Nylstroom): between Groenvallei and Roed-

tan, Farm Zoetkoppies (-BB), S.M. Perold 339 p.p. (PRE). 2725

(Bloemhof): Farm Leeufontein, 10 km W of Wolmaransstad

(-BB), A.E. van Wyk 5753 p.p. (PRE).

O.F.S. — 2825 (Boshof): Farm Goede Hoop (-CA), Volk

81-204 p.p., 81-210 p.p. (M, PRE). 2827 (Senekal): 5 km S of

turnoff to Willem Pretorius Wildtuin (-AC), S.M. Perold 957

(PRE). 2926 (Bloemfontein): nr Eagle’s Nest, Bloemfontein

(-AA), Potts 7003B-E (BLFU).

CAPE. — 2922 (Prieska): Farm Erfrust, N bank of Orange

River, between Prieska and Koegasbrug (-BC), Smook 4487

(PRE).

FIGURE 5. — Distribution map of R. argenteolimbata, ■; and

R. albomata, 0, in southern Africa.

2. Riccia albomata Volk & Perold, sp. nov.

Thallus monoicus, mediocris, viridis; in sicco albidus,

nitidus. From usque ad 12 mm longa, 1,5— 2,0(-4,0)

mm lata, 1,0— 2,0 mm crassa, triplo latior quam crassa,

bi- vel trifurcata; lobis terminalibus brevibus, valde di-

vergentibus, late ligulatis, apice rotundato emarginato, in

partibus junioribus profunde acuteque sulcatis; cellulae

dorsales epithelii grandes, ad 60 pm diametro, saepe duas

columnas chlorophylloferas obtegentes; chlorenchyma

dimidio crassitudinis thalli, canalae aeriferae ad 80 /urn

latae ab 6 columnis chlorophyllosis circumcinctae; costa

subplana, lateribus oblique in margines acutes excurren-

tibus, viridibus vel purpureis. Squamae magnae, crispatae,

calce incrustatae, vel hyalinae, marginem frondis exce-

dentes, dense imbricatae, basi saepe coloratae. Sporae

polares, ± 100 pm diametro, alatae, ± irregulariter reti-

culatae, foveolis 2,5 pm latis, saepe imperfectis. Ricciae

albolimbatae similis, sed ornamentatione sporarum et

conformatione epithelii et chlorenchymatis differt.

Chromosomatum numerus n = 15.

TYPE. — Cape Province, 2921 (Kenhardt): ± 10 km

westl. Kenhardt, an der Strasse nach Kakamas (—AC),

Bothalia 18,2 (1988)

alt. 970 m, grobsandiger, schluffreicher, rotbrauner Gra-

nitzersatz am Fuss von Batholithen, pH 7,1, mitR. oka-

handjana S. Amell undMoosen, 1981.01.15, Volk 81-081

(M, holo.).

Thallus monoicous (Figures 6A, 7B), perennial, in

crowded gregarious patches, not in rosettes; branches sim-

ple or bi- to trifurcate, symmetric or asymmetric, usually

widely divergent, oblong, (4— )5—9(— 12) mm long, 1,5—

2,0(-4,0) mm broad, 1,0— 2,0 mm thick, 1,2— 2,5 or

more times broader than thick; apex rounded, obtuse,

emarginate, sulcus deep apically (Figures 6A; 7A, B, C),

gradually becoming shallow and wide, flattening out

proximally (Figure 6B1— 4); dorsally green, shiny, older

surfaces and edges yellowish or whitish green; margins

acute, slightly attenuate, flanks sloping obliquely outward,

violet or green; ventral surface flat to rounded, green;

when dry, greyish white and margins inflexed with large,

frilly, hyaline or lime-encrusted, white scales covering

most of dorsal surface.

Anatomy of thallus : dorsal epithelial cells globose or

mammillate (Figures 6C; 7C, D), 30-40(-50) x 40-60

pm, often spanning two columns of assimilation cells be-

neath (Figure 6C, D, centre), surface occasionally lightly

dusted with fine calcium carbonate deposits (Figure 7F),

towards margins and in older parts cells frequently col-

lapsed (Figure 7E); air pores narrowly rectangular or

wider near the margins and 5 or 6-sided, positioned over

wide, mostly 6-sided air canals, up to 80 pm wide (Figure

6D); assimilation tissue (chlorenchyma) about V2 the

thickness of thallus, cells short-rectangular, 40—50 x ±

30 pm wide, in columns of 6— 8(— 10) cells; storage tissue

about \ the thickness of thallus, cells ± 50 jum wide.

Rhizoids mostly smooth, some tuberculate, ± 20 pm

wide. Scales imbricate, rounded, margin smooth, project-

ing ± 50—100 pm above margin of thallus, those at apex

hyaline, more proximal ones white, sometimes with red-

dish purple bases, large, up to 1 250 x 750 pm, cells oc-

casionally lime-encrusted, walls thick, 4— 6-sided, 75—85

(—100) x 35—50 pm in body of scale; brick-shaped, ±

35 x 60 pm wide (Figure 6E) in margin. Antheridia with

prominent hyaline necks projecting along midline. Arche-

gonia with purple necks scattered singly along either side

of midline. Sporangia containing ±300 spores each, cover-

ing tissue with large pores, disintegrating when ripe and

leaving capsules exposed in hollow. Spores large, (85—)

95— 105(— 115) pm in diameter, straw-coloured or yel-

low to brown, semi-transparent to opaque; triangular-

globular, polar, with wing up to 5 pm wide (Figure 8A,

B), notched or perforated at angles, margin finely crenu-

late, distal face ± reticulate with 14 — 16(— 20) small, deep

areolae across the diameter, about 2,5 pm wide (Figure

8C, D, F); areolar walls ± 2,5 pm thick, raised into pro-

cesses at the nodes (Figure 8E) and frequently anasto-

mosing to form short convoluted ridges (Figure 8C, D);

proximal face with triradiate mark distinct, about 30— 40

small areolae on each of 3 facets (Figure 8A). Chromo-

some number n = 1 5 (Figure 6F).

This unusual haploid set with 1 5 chromosomes (Figure

6F) is known only in R. albomata ( Volk 81-081 — 15

counts); Volk 84-667 p.p. — 4 counts). The chromo-

some formula of this set contains, in addition to the basic

set, 2B, , 2C| and 3D, chromosomes.

Vegetatively R. albomata is not easily distinguished

from R. albolimbata, but it never grows in rosettes, the

Bothalia 18,2 (1988)

161

FIGURE 6. — Riccia albomata. Morphology and anatomy. A, fresh thalli with sporangia, archegonial and antheridial necks; Bl— 4,

transverse sections of branch at different distances from apex to proximal part; C, enlargement of transverse section through

dorsal epithelium (showing wide cells) and through assimilation tissue; D, horizontal section at different levels: left, air

pores and epithelial cells; in centre, position of air pores over air canals with epithelial cells extending over columns of

chlorenchyma; right, chlorenchyma with air canals; E, scale; F, chromosomes. (A-E by Volk; F by Bomefeld) (A-D, F,

Volk 81-081; E, Volk 84-667). Scale bars: A, B = 1 mm; C, D =50 |um; E =100 um; F = 1 Mm.

162

Bothalia 18,2 (1988)

FIGURE 7. — Riccia albomata.

Morphology and anatomy.

A, dorsal view of apex and

groove with partly inflexed

margins and large scales; B,

archegonial and antheridial

necks; C, dorsal cells in

groove; D, more enlarged

view of dorsal cells at

groove; E, dorsal cells and

air pores toward margin;

F, dorsal cells with deposits

(SEM micrographs by Pe-

rold). (A, B, E, F, Oliver

8854a; C, D, Morley

CH4525.) Scale bars on

A— F = 100 Mm.

hyaline scales are larger and frillier, the dorsal epithelial

cells are also larger, the air canals wider and generally en-

closed by six columns of cells and the spore ornamentation

is markedly different with numerous small, irregular

areolae and convoluted ridges on both faces. The distri-

bution areas of the two species do not appear to overlap

either, except for two localities in the central Cape.

R. argenteolimbata is generally a smaller plant, with

regular, stiff white scales, a mat, glaucous grey dorsal

surface and apolar spores; it is also dioicous.

R. albomata is a rare endemic species infrequently

collected at altitudes of 500—1 500 m above sea level. It

appears to be confined to those regions of the Cape Pro-

vince (Figure 5) which are influenced by, or generally

receive sparse winter rains of at least 125 mm per annum.

Generally it grows on reddish brown or light brown,

coarse, gravelly soil, derived from granite or quartzite,

on koppies or on slopes, in full sun or in light shade of

small shrubs or thorn trees. The soil pH values are 7,0—

8,0 (Volk). It is sometimes associated with other Riccia

spp., e.g. R. okahandjana S. Arnell and with mosses, e.g.

Desmatodon convolutus (Brid.) Grout and Funaria spp.

SPECIMENS EXAMINED

CAPE. — 2820 (Kakamas): Aughrabies Nat. Park, Oranje-

kom (-CB), Volk 84-667 (M, PRE). 2917 (Hondeklipbaai):

36 km S of Springbok, on road to Kamieskroon (-DD), S.M.

Perold 1445 (PRE). 2921 (Kenhardt): 10 km W of Kenhardt

(-AC), Volk 81-081 (M, PRE). 3119 (Calvinia): E of Slagberg,

midway between Nieuwoudtville and Loeriesfontein, Farm

Koringhuis (-AB), S.M. Perold 1800, 1801 (PRE); NE of

Nieuwoudtville, Groothoek, on Soetlandsrivier (-AD), Oliver

8854a (PRE). 3123 (Victoria West): Farm Rietpoort, 34 km N

of Victoria West, on slope (-AA), Smook 6961 (PRE); Farm

Kalkfontein, 48 km NE of Victoria West, on flat gravel plain

(-AA), Smook 6990 (PRE). 3125 (Steynsburg): White Ridge

(-AC), Duthie 5149 (BOL). 3218 (Clanwilliam): Bidouw, Farm

Mertenhof (-BB), Oliver 1463 (BOL). 3222 (Beaufort West):

Beaufort West, on small koppie (-BC), Steyn 5487 (BOL). 3319

(Worcester): Rabiesberg, Farm Doringkloof (-DA), Morley

CH4525 (PRE); Nuy (-DA), Morley 362 (PRE). 3420 (Bredas-

dorp): De Hoop, in pass on road from Wydgelee to De Hoop

(-AD), Fellingham 746a (PRE).

ACKNOWLEDGEMENTS

Sincere thanks are due to the Curators of Botanische

Staatssammlung, Munchen and Bolus Herbarium, Uni-

versity of Cape Town, for the loan of specimens and to

the many people who kindly collected specimens for us,

especially Botanical Research Institute staff members,

Dr G.E. Gibbs Russell, Messrs D. Hardy, E.G.H. Oliver,

Mrs L. Fish (nee Smook), Misses E. Retief, L. Hender-

son and M. Morley as well as to the Hoffmann family,

Farm Oros, SWA/Namibia.

Bothalia 18,2 (1988)

163

FIGURE 8. — Riccia albomata.

Spores. A, proximal face;

B, side view of proximal

face; C, D, F, distal face;

E, more enlarged view of

areolae near margin on

distal side. (A-E, SEM

micrographs; F, LM (light

microscope) micrograph by

Perold.) (A-D , Oliver 5149;

E, F, Volk 84-667 .) Scale

bars on A-E =50 /am; dia-

meter of spore on F, ± 100

/am.

REFERENCES

ARNELL, S. 1957. Hepaticae collected in South West Africa

by Prof. Dr O.H. Volk. Mitteilungen ms der botanischen

Staatssammlung, Miinchen 16: 262-272.

ARNELL, S. 1963 Hepaticae of South Africa, pp. 411. Swedish

National Scientific Research Council, Stockholm.

BORNEFELD, T. 1984. Chromosomenanalyse der Gattung Ric-

cia von SUd- und SW-Afrika und allgemeine Bemerkungen

zur Zytogenetik der Lebermoose. Nova Hedwigia 40:

313-328.

BORNEFELD, T. 1987. The natural system of the Marchantiales

based upon cytogenetical and morphological evidence.

Nova Hedwigia 45 : 41-52.

BORNEFELD, T. & GRILLENBERGER, Chr. 1987. Some as-

pects of size in chromosome analysis of liverworts (Mar-

chantiales). Nova Hedwigia 44: 91-100.

PEROLD, S.M. in press. Studies in the genus Riccia (Marchantiales)

from southern Africa. 12. Riccia albolimbata and the

status of R. albosquamata. Bothalia.

VOLK, O.H. 1966. Die Florengebiete von Siidwestafrika. Journal

SWA Wissenschaftliche Gesellschaft Windhoek 20: 25-58.

VOLK, O.H. 1983. Vorschlag fur eine Neugliederung der Gattung

Riccia L. Mitteilungen ms der botanischen Staatssamm-

lung, Miinchen 19: 453—465.

VOLK, O.H. 1984. Pflanzenvergesellschaftungen mit Riccia- Allan

in Siidwestafrika (Namibia). Vegetatio 55: 57-64.

VOLK, O.H. 1984. Beitrage zur Kenntnis der Marchantiales in

Siidwestafrika (Namibia). Nova Hedwigia 39: 117-143.

Bothalia 18,2 : 165-171 (1988)

Taxonomy and leaf anatomy of the genus Ehrharta (Poaceae) in

southern Africa: the Dura group

G.E. GIBBS RUSSELL* and R.P. ELLIS*

Keywords: Capensis, Ehrharta, leaf anatomy, Mountain Fynbos, Poaceae, taxonomy

ABSTRACT

The Dura species group in the genus Ehrharta Thunb. is differentiated morphologically by the perennial habit

and the very large, awned, subglabrous spikelets and anatomically by the occurrence of tanniniferous cells and

wax platelets obscuring the stomatal pores. The Dura group consists of two species, E. dura Nees ex Trim and E.

microlaena Nees ex Trim, which occur only in Mountain Fynbos. The group shows no clear morphological or

anatomical relationship with other species groups in the genus in southern Africa.

UITTREKSEL

Die Dura-spesiegroep in die genus Ehrharta Thunb. word morfologies deur die meerjarige groeiwyse en die

besonder groot, genaalde, ylbehaarde blompakkies onderskei en anatomies deur die aanwesigheid van tannien-

selle en wasplaatjies wat die huidmondjieporiee verberg. Die Dura-groep bestaan uit twee spesies, E. dura Nees

ex Trim en E. microlaena Nees ex Trim, wat slegs in Bergfynbos voorkom. Die groep vertoon geen duidelike mor-

fologiese of anatomiese verwantskap met ander spesiegroepe in die genus in suidelike Afrika nie.

INTRODUCTION

Previous papers in this series have outlined the seven

provisional species groups for Ehrharta in southern Afri-

ca, and dealt with the Setacea and Villosa groups (Gibbs

Russell & Ellis 1987; Ellis 1987a, 1987b; Gibbs Russell

1987a, 1987b). The Dura group is a small, apparently

isolated group composed of only two species, E. dura

Nees ex Trin. and E. microlaena Nees ex Trim The two

species are distinguished morphologically from other

Ehrharta species by correlation of perennial habit, large

spikelets 9—17 mm long and awned sterile lemmas with

glabrous sides, keel and margins. The spikelet plan of the

Dura group is shown in Figure 1 . Anatomically the Dura

group is distinguished within Ehrharta by the presence

of tanniniferous cells and by wax platelets obscuring the

stomatal pores. The spikelets and leaves of the species

have a characteristic olive-green colour that is distinctive

and may result from the dark tannins in the intercostal

long cells of the epidermis. The species differ from each

other principally in awn length and in leaf blade shape

and width. Both species occur only in Mountain Fynbos.

The two species in the group present no nomenclatural

difficulties because they were described by Trinius (1839)

in the same publication, and other authors have consis-

tently followed this treatment (Nees 1841; Steudel

1853; Stapf 1900; Chippindall 1955); thus neither

species has synonyms. The same treatment is followed

here. This singularly simple situation is undoubtedly a

reflection of the distinctness of the Dura group within

Ehrharta and the unusually low degree of intraspecific

variation present. It is more difficult to assess to what

degree the simplicity of treatment also results from the

infrequent collection of the species due to their restrict-

* Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

MS. received: 1987.12.02.

ed and specialized distribution and habitat and the fact

that they are apparent in vegetation only after fire.

However, lectotypification of the names is necessary.

Trinius (1839) published descriptions of both species in

his treatment of the tribe Phalarideae without citing any

specimens. However, Trinius ascribed both names to

Nees, presumably because he had obtained specimens

from Nees with manuscript names. Two years later, Nees

(1841) also published descriptions, for which he claimed

authorship, in his treatment of all southern African gras-

ses. Specimens of both species clearly labelled in Nees’

handwriting exist in several herbaria. The Drege specimen

of each species in the Herbarium, Komarov Botanical

Institute, Leningrad (LE) is designated as lectotype, be-

cause these specimens were seen by Trinius who first

published a description.

Two previous studies, Engelbrecht (1956) and Tateoka

(1963), have covered some aspects of leaf anatomy of

this group, and their findings are in accord with the de-

tailed results presented here.

FIGURE 1. — Spikelet of E. dura ( Ellis 5457, PRE): a,

whole spikelet; b, glumes; c, first sterile lemma;

d, lemma of fertile floret; e, palea of fertile floret;

f, second sterile lemma; all X 2.

166

METHODS

Methods adopted for previous papers in this series

were followed here, both for the taxonomic and the

anatomical studies. The descriptions and keys were pre-

pared through the DELTA computer system for handling

descriptive data (Dallwitz 1984).

TAXONOMY

Key to species

Leaves with blades expanded, 4-10 mm across, lanceolate;

basal sheaths persistent, hard, reddish brown; awns

2-16 mm long E. dura

Leaves with blades reduced, setaceous, to 1 mm across;

basal sheaths eventually deciduous, membranous,

light brown or whitish; awns 13-25 mm long . . .

E. microlaena

1. Ehrharta dura Nees ex Trin. in Phalaridea, Me-

moires de l’Academie imperial des Sciences de St-Peters-

burg, ser. 6, 5: 13 (of reprint) (1839); Nees: 218(1841);

Steudel: 5 (1853); Stapf: 665 (1900); Chippindall: 37

(1955); Smook & Gibbs Russell: 55 (1985). Type:

Drege, Du Toit’s Kloof, 3 000 ft, (LE!, lecto., here de-

signated; K!,P!).

Perennial, erect, forming large leafy tufts, rarely long-

rhizomatous. Rhizomes woody, with glabrous, thickened

cataphylls. Culms several to many, to 800 mm long, 4

mm across, herbaceous, crowded, unbranched, hollow.

Young shoots intravaginal. Leaves mostly basal, culm

leaves with blades well developed, sheaths not overlap-

ping; leaves auriculate, auricles produced from top of

sheath; basal sheaths flabellate and slightly twisted,

tight, persistent, hard, reddish brown, bearing blades;

ligule a membrane fringed with hairs or sometimes near-

ly glabrous, 1, 5-3,0 mm long; leaf blades persistent,

erect, herbaceous, glabrous, lanceolate, 4—10 mm across,

flat or folded, becoming rolled from margins when

dry; blade bases folded, thickening with age, persistent

and conspicuous often even in burned plants; blade tips

gradually tapering. Inflorescence a panicle, open, 70—200

mm long, considerably overtopping leaves; main axis

straight, often closely subtended or enveloped below by

uppermost leaf sheath; numerous spikelets, spikelets

appressed to the axes and held nearly erect but somewhat

secund. Spikelets pedicellate, distinctly compressed

laterally, 9—16 mm long, 3 mm across laterally (above

glumes), olive-green. Rhachilla prolonged beyond

fertile floret. Glumes not keeled, the lower V2 or more

the length of the upper, V4-V2 as long as spikelet, white,

or green, appressed to lemmas at maturity. Lower glume

2,5— 6,5 mm long, 3— 5-nerved, acute. Upper glume 3,5—

7,0 mm long, 5— 9-nerved, truncate apically. Florets

with lemmas decidedly firmer than the glumes, keeled.

Sterile lemmas laterally compressed, sides flat, similar in

shape, texture and ornamentation; with keel and margins

parallel; base not stipitate, without auriculate appendages,

shortly bearded, sides scabrous or sometimes with short

hairs, dull, with 3—1 1 strong or fairly distinct longitudi-

nal ribs; tip tapering gradually from body of lemma to

an arista or awn. Awn 2—16 mm long. First sterile lem-

ma about V2 to z/3 length of second sterile lemma. Fer-

tile floret shorter than second sterile lemma. Lemma of

fertile floret differing from sterile lemmas, strongly

Bothalia 18,2 (1988)

laterally compressed and sides unomamented, faintly

3— 5-nerved, sides glabrous, tip truncate with a tuft of

minute hairs. Palea thinner than lemma, 3/4 or more as

long as lemma, keeled, 1-nerved. Lodicules 2, membra-

nous, 2-lobed, margins ciliate. Stamens 4 or 6. Anthers

7 mm long, yellow. Ovary glabrous. Stigmas white.

Caryopsis 9 mm long, laterally flattened.

E. dura is distinguished from southern African Ehr-

harta species in other groups by the perennial habit and

the large nearly glabrous mucronate to awned spikelets.

It is distinguished from closely related E. microlaena by

the broad leaves and the broad, brown, flabellate bases.

It grows in mesophytic to seasonally moist open habitats

in Mountain Fynbos in sandy or loam soils over sand-

stone or granite at altitudes of 430 to 1 300 m. A single

specimen ( Boucher 2028) came from an altitude of 30

m at Kogelberg State Forest. All but a few of the speci-

mens seen were collected after a bum. E. dura extends

along mountain ranges from Stellenbosch and Worcester

to Humansdorp (Figure 2), and has a much wider distri-

bution than E. microlaena. Most Ehrharta species show

considerable variation in habit and leaf development,

and it is possible to recognize ecotypes and local variants.

In contrast to this common pattern, E. dura is remark-

ably constant in its appearance from the eastern to the

western ends of its range, with no geographically corre-

lated infraspecific variation. Flowering occurs from

September to December.

Vouchers: Acocks 22817; Bond 1611; Taylor 4211, 10256;

Zeyher 4513.

2. Ehrharta microlaena Afees ex Trin. in Phalaridea,

Memoires de l’Academie imperial des Sciences de St-

Petersbourg, ser. 6, 5: 13 (of reprint) (1939); Nees: 218

(1841); Steudel: 5 (1853); Stapf: 665 (1900); Chip-

pindall: 37 (1955); Smook & Gibbs Russell: 55 (1985).

Type: Drege, Du Toits’ Kloof (LE!, lecto., here designa-

ted; K!, SAM!).

Perennial, erect, tufted, apparently lacking long rhi-

zomes. Culms several, to 1 100 mm long, 2,5 mm across,

herbaceous, crowded, unbranched, hollow. Young shoots

intravaginal. Leaves mostly basal, culm leaves with blades

reduced, sheaths not overlapping. Leaves auriculate, the

auricles produced from the base of the blade and adnate

to the margins of the ligule. Basal sheaths somewhat

flabellate, tight, eventually deciduous, membranous,

light brown, or whitish, bearing blades. Ligule a glabrous

Bothalia 18,2 (1988)

membrane to 8 mm long. Leaf blades persistent, erect,

curved slightly outward from middle, herbaceous,

glabrous, linear, reduced, setaceous, to 1 mm across;

blade tips gradually tapering. Inflorescence a panicle,

open, or somewhat contracted, 6—18 mm long, con-

siderably overtopping leaves; main axis straight, usually

exserted from uppermost leaf sheath; of numerous

spikelets, spikelets spreading and somewhat secund.

Spikelets pedicellate, distinctly compressed laterally,

13—15 mm long, to 2,5 mm across laterally (above

glumes), with conventional internode spacing. Rachilla

prolonged beyond fertile floret. Glumes not keeled,

the lower V2 or more the length of the upper, V4-V3 as

long as spikelet, green, appressed to lemmas at maturity.

Lower glume 2,5— 3,0 mm long, 3-nerved, acute. Upper

glume 4—5 mm long, 7-nerved, truncate apically. Florets

with lemmas decidedly firmer than the glumes, keeled.

Sterile lemmas laterally compressed, sides flat, similar

in shape, texture and ornamentation; with keel and

margins parallel; base not stipitate, without auriculate

appendages, sparsely and shortly bearded at base; sides

glabrous or puberulous, dull, with 5—7 longitudinal ribs;

tip tapering gradually from body of lemma into a long

awn. Awn as long as or longer than body of lemma, 13—

25 mm long. First sterile lemma about \ length of

second sterile lemma. Fertile floret shorter than second

sterile lemma. Lemma of fertile floret differing from

sterile lemmas, strongly laterally compressed and sides

unornamented, obscurely 5— 7-nerved, sides glabrous,

tip truncate with a tuft of minute hairs. Palea thinner

than lemma, 1 * 3/4 or more as long as lemma, keeled, 1-

nerved. Lodicules 2, membranous, 2-lobed, margins

ciliate and laciniate. Stamens 4. Anthers 7 mm long,

yellow. Ovary glabrous. Stigmas white. Caryopsis 8 mm

long, laterally flattened.

E. microlaena is easily distinguished from all other

Ehrharta species by the perennial habit, the large, glabrous,

long-awned spikelets and the setaceous leaf blades. It is

hydrophytic, and grows on streamsides and in damp peaty

places in Mountain Fynbos. Like E. dura , most of the

specimens seen were collected after fire. However, the

distribution of E. microlaena is more restricted than that

of E. dura, extending from the Kogelberg Forest Reserve

north through Paarl, Bain’s Kloof, Du Toit’s Kltiof and

Ceres to Tulbagh (Figure 3) at altitudes of 400 to 1 330

m. There is little intraspecific variation, and specimens

from throughout the range are similar in appearance.

Flowering occurs from December to February.

Vouchers: Adamson 3603; Boucher 173, 1838; Esterhuy sen

28427, 35600.

1. LEAF ANATOMY OF E. DURA

Transverse section

The leaf blade is open and expanded (Figure 4A) but

has the ability to inroll from the margins under adverse

moisture conditions. The midrib is not structurally dis-

tinguishable from the lateral first order vascular bundles

and is not associated with colourless parenchyma. Suc-

cessive first order bundles are separated by three smaller

bundles, usually two third order bundles with a single

second order bundle between them (Figure 4A) although

this pattern is not always consistent (Figure 4B).

167

Well developed, rounded to slightly flattened adaxial

ribs are located over all the vascular bundles (Figure 4A—

C). These ribs generally are all the same size but the

smallest third order bundles may be associated with

smaller ribs (Figure 4B). The furrows between adjacent

ribs are deep, narrow and steep-sided and tend to become

cleft-like with leaf inrolling. Conspicuous bulliform cells

occur at the bottom of these furrows (Figure 4C).

The mesophyll tissue is irregularly arranged. The chlo-

renchyma cells are larger than the bundle sheath cells,

angular and fitting tightly together with the chloroplasts

concentrated around the cell circumference (Figure 4C).

The walls are straight with no arm cell-like invaginations.

No large intercellular air spaces are visible. Colourless

parenchyma is absent between the bundles although

colourless adaxial bundle sheath extensions are associated

with the smaller bundles (Figure 4B, C). The vascular

bundles are surrounded by two bundle sheaths, an outer

sheath of many (up to 26) small colourless cells and a

distinct mestome sheath of cells with uniformly thicken-

ed secondary walls (Figure 4C). All the epidermal cells

contain conspicuous, dark, resin-like deposits distributed

evenly across the entire width of the blade (Figure 4B,

C).

Abaxial epidermis

Costal and intercostal zones are differentiated (Figure

4D, E). The intercostal long cells are rectangular with

slightly wavy anticlinal walls (Figure 4D, E) and inflated

outer walls (Figure 5A— C). All contain black tanninife-

rous deposits generally filling the entire cell lumen (Figure

4E). Adjacent long cells in a file are separated by single

or paired short cells (Figure 4D, E). Cell size and arrange-

ment is uniform across each intercostal zone.

Stomata occur in a single file on either side of each

intercostal zone, immediately adjacent to the costal

zones (Figure 4E). Successive stomata are separated by

single interstomatals and neither the subsidiary cells

nor the interstomatals contain tanniniferous deposits.

The subsidiary cells are low dome-shaped and not sunken

below the general level of the epidermis (Figure 5C, D).

The stomatal pores are obscured by dense accumulations

of wax platelets (Figure 5D).

The costal silica bodies are irregularly dumbbell-shaped

and alternate with similarly shaped short cells. Small

prickle-hairs with very short barbs are interspaced at

168

Bothalia 18,2 (1988)

FIGURE 4. — Leaf anatomy of Ehrharta dura. A-C, leaf in transverse section, Ellis 4695: A, outline showing absence of keel,

scale = 20 jum; B, prominent adaxial ribs and deep furrows, scale = 10 (im; C, detail of chlorenchyma and tanniniferous

epidermal cells (t), scale = 5 jum. D-E, abaxial epidermis: D, Ellis 4695, epidermal zonation and tanniniferous intercostal

long cells (t), scale = 10 Atm; E, Ellis 4696, all intercostal long cells with tanniniferous deposits (t), scale = 10 Atm.

intervals along the costal zones (Figures 4D, E). Micro-

hairs were not observed with the light microscope but

are clearly visible with the SEM (Figure 5C, E). These

hairs have a short basal cell with a blunt distal cell

(Figure 5E). Small pointed, single-celled intercostal

macrohairs are very rare (Figure 5F).

Specimens examined

CAPE. — 3318 (Cape Town): Stellenbosch (-DD), Taylor

10256. 3320 (Montagu): Grootvadersbos (-DD), Ellis 5554.

3321 (Ladismith): Cloetes Pass, Langeberge (-DD), Ellis 4695,

4696. 3322 (Oudtshoorn): Robinson Pass, Outeniqua Mts

(-CC), Ellis 5457. 3324 (Steytlerville): Groot Winterhoek Mts,

Cockscomb Peak (-DB), Ellis 5615. 3419 (Caledon): Rivier-

sonderend Mts, Die Galg (—BA), Ellis 5568.

2. LEAF ANATOMY OF E. MICROLAENA

Transverse section

No fresh leaf blade material of this taxon was avail-

able for anatomical study, and consequently the transec-

tional anatomy was not examined. Engelbrecht (1956)

has shown that the leaf blade is cylindrical in section

with the interior containing colourless parenchyma tissue.

This structure appears to result from the fusion of the

two margins and not from a reduction of the lamina on

either side of the midrib with the resultant blade being

only a rounded keel. This specialized blade needs detailed

anatomical study as it represents an interesting adaptation

within the genus.

Abaxial epidermis

Herbarium leaf blade material was used to prepare

abaxial epidermal scrapes of this species. The epidermal

structure is very similar to that of E. dura, and reference

should be made to the description above. Both species

have the diagnostic tanniniferous intercostal long cells

which dominate the epidermis and are unique to this

species group in Ehrharta.

Specimens examined

CAPE. — 3319 (Worcester): Hex River Mts, Milner Peak

(-AD), Ellis 5524\ Du Toit’s Kloof (-CA), Boucher 1838.

DISCUSSION AND CONCLUSIONS

The two species in the Dura group are the only awned

perennial Ehrharta species in southern Africa. Three

other species have awns, but all are annual: E. longiflora

and E. triandra in the Erecta group and E. pusilla in the

closely related Calycina group. These three species grow

near ephemeral water bodies, and each is apparently

most closely related to the unawned perennial species

within their group. Awns are therefore probably the

result of parallel or convergent evolution in different

species groups. Therefore, no phylogenetic relationship

between the Dura group and the annual awned members

of the Erecta and Calycina groups should be inferred on

the basis of awn occurrence.

Bothalia 18,2 (1988)

169

The spikelet size of the Dura group species, excluding

the awns, is only equalled in the three taxa of the Villosa

group: E. thunbergii, E. villosa var. villosa and E. villosa

var. maxima (Gibbs Russell 1987b). However, the shape

and vesture of the sterile lemmas differ and are distinc-

tive for both groups, and size alone is poor evidence upon

which to speculate relationship.

The spikelets of the Dura group are distinctive in

other respects as well, and therefore the Dura group

cannot be shown to have a close similarity with any

other species group on the basis of spikelet structure

(Gibbs Russell & Ellis 1987). However, the consistent

occurrence of four stamens in E. microlaena and four or

six stamens (the usual number for southern African

Ehrharta species) in E. dura, suggests that relationships

for the Dura group might be sought within the Austra-

lian species formerly assigned to Tetrarrhena (Willemse

1982). Tetrarrhena is characterized by the presence of

four stamens, a most unusual condition in the Poaceae.

The Dura species group has distinctive leaf anatomy

that separates it from the other groups in the genus in

southern Africa (Gibbs Russell & Ellis 1987). In transec-

tion the leaf blade lacks a keel, has well developed, flat-

tened adaxial ribs and furrows and the mesophyll consists

of large angular chlorenchyma cells with no definite pat-

tern of arrangement. The tanniniferous epidermal cells,

particularly the bulliform cells, are diagnostic for this

group. This anatomy superficially resembles the Villosa

group but the two groups can be separated by the flat

ribs and tanniniferous cells of the Dura group and the

FIGURE 5. — Ultrastructure of the abaxial epidermis of E. dura. A, Ellis 4695 with inflated, thick-walled intercostal long cells,

stomatal file and costal zone with row of prickles, X 200; B, Ellis 4696 showing intercostal long cells, stomata and a prickle,

X 100; C-E, Ellis 4695: C, more intercostal long cell detail with microhairs and stomata visible, X 150; D, stoma with

dense accumulation of wax platelets overlying the pore, X 500; E, microhair with short basal cell and longer blunt distal

cell, X 500; F, Ellis 4696 showing small intercostal macrohair, X 500.

170

tendency towards semi-radiate chlorenchyma with

abaxial palisade-like cells in the Villosa group (Ellis

1987). The Setacea (Ellis 1987) and Ramosa groups and

the Longifolia subgroup all lack keels but differ from the

Dura group in many other respects (Gibbs Russell &

Ellis 1987).

The abaxial epidermis is differentiated into costal and

intercostal zones with rectangular long cells, low dome-

shaped subsidiary cells, irregularly dumbbell-shaped

silica bodies, prickles with short barbs and microhairs

with blunt distal cells. Epicuticular wax is absent except

in association with the stomatal apertures where wax

platelets obscure the pores. This combination of epider-

mal attributes is distinctive and serves to distinguish the

Dura group from the others (Gibbs Russell & Ellis 1987).

Rectangular long cells are shared with the Villosa and

Ramosa groups but the stomata, silica bodies and micro-

hairs differ between these three groups.

The unique tanniniferous epidermal cells found in the

species of the Dura group are of great interest because

these represent the only known C3 grasses which possess

this type of cell (Ellis in prep.). No bambusoids, pooids

or other arundinoids are known to have this type of

epidermal cell which is also rare in the chloridoids and

panicoids, except the Andropogoneae and Arundinelleae,

where many species and genera exhibit tanniniferous

cells. The colour and texture of the contents of the

epidermal cells of E. dura and E. microlaena are con-

sistent with that of the tanniniferous cells observed in

these C4 grass taxa and it is presumed that they contain

the same chemical substance. All 90 South African grass

species in which this type of cell has been observed are

unpalatable species from oligotrophic soils and it is in-

ferred that these cellular contents are polyphenols which

function as a chemical defence against herbivores (Ellis

in prep.). Their presence in the Dura group may repre-

sent a separate and similar adaptation to which no phy-

logenetic importance should be attached.

The anatomical observations of this study generally

agree with previous reports but for a minor exception.

Tateoka (1963) described the subsidiary cells of if. dura

as being almost parallel-sided, whereas here they are

shown to be low dome-shaped. However, the distinction

between these two stomatal types is only slight and a

difference in interpretation may have resulted in this

apparent inconsistency. The work of Engelbrecht (1956)

is in full agreement with this study.

It is of interest that in one of the five specimens of

E. dura examined by Engelbrecht (1956) a distinct keel,

incorporating five vascular bundles with extensive adaxial

parenchyma, was observed. This appears to be an inter-

mediate condition between E. dura and the cylindrical

blade of E. microlaena. It appears as if the cylindrical

leaf blade of E. microlaena represents an adaptation to

hygrophilous habitats and the intermediate specimen

probably came from a damp locality. Unfortunately it

is not possible to deduce which specimen displayed this

anatomy from the work of Engelbrecht (1956). Never-

theless, this represents the first possible intermediate

between these two species which are otherwise so

clearly separated anatomically and morphologically.

E. dura and E. microlaena are undoubtedly closely

related to each other, yet have very little in comrpon

Bothalia 18,2 (1988)

with any other species of Ehrharta in southern Africa.

Both spikelet morphology and leaf anatomy indicate

that they occupy an isolated position within the genus

and are not linked to any of the other species groups.

The two taxa are treated at the level of species because

they are differentiated by both vegetative and spikelet

characteristics, have very different transectional leaf

blade anatomy and their habitats are somewhat different.

In addition, although the two taxa occur sympatrically,

they flower at different times and no morphologically

intermediate specimens have been observed.

ACKNOWLEDGEMENTS

We thank the Directors of the Herbaria that have lent

specimens previously: BOL, J, JF, K, NBG, SAM, STE,

and in addition, P and LE, as well as the Director of B

for providing visitors’ facilities for study of a number of

grass genera. We thank G. Condy for the spikelet drawing,

S.Perold for scanning electron microscopy, A. Romanow-

ski for photography, H.Ebertsohn for technical assistance

and W. Roux for technical assistance and map preparation.

REFERENCES

CHIPPINDALL, L. 1955. A guide to the identification of grasses

in South Africa. In C. Meredith, The grasses and pastures

of South Africa. Central News Agency, Cape Town.

DALLWITZ, M.J. 1984. User’s guide to the DELTA system — a

genera system for coding taxonomic descriptions. CSIRO

Division of Entomology Report No. 13.

ELLIS, R.P. 1987a. Leaf anatomy of the genus Ehrharta (Po-

aceae): the Setacea group. Bothalia 17: 75-89.

ELLIS, R.P. 1987b. Leaf anatomy of the genus Ehrharta (Po-

aceae): the Villosa group. Bothalia 17: 195-204.

ENGELBRECHT, A.H.P. 1956. 'n Morfologiese studie van die

genus Ehrharta Thunb. M.Sc. thesis, University of Pretoria.

GIBBS RUSSELL, G.E. 1987a. Taxonomy of the genus Ehrharta

(Poaceae): the Setacea group. Bothalia 17: 63-73.

GIBBS RUSSELL, G.E. 1987b. Taxonomy of the genus Ehrharta

(Poaceae): the Villosa group. Bothalia 17: 191-194.

GIBBS RUSSELL, G.E. & ELLIS, R.P. 1987. Species groups in

the genus Ehrharta (Poaceae) in southern Africa. Bothalia

17: 51-65.

NEES AB ESENBECK, G.C. 1841. Florae Africae Australioris.

Prausnitzianus, Cracow.

SMOOK, L. & GIBBS RUSSELL, G.E. 1985. Poaceae. In G.E.

Gibbs Russell et al., List of species of southern African

plants, edn 2, part 1: 45-70. Memoirs of the Botanical

Survey of South Africa No. 51.

STAPF, O. 1900. Gramineae. In W.T. Thiselton-Dyer, Flora

capensis 8: 310-750.

STEUDEL, E.G. 1853. Synopsis plantarum graminearum . Metzler,

Stuttgart.

TATEOKA, T. 1963. Notes on some grasses. XII. Relationships

between Oryzeae and Ehrharteae, with special reference

to leaf anatomy and histology. The Botanical Gazette 124:

264-270.

TRINIUS, C.B. 1839. Phalaridea. Memoires del Academie impe-

rial des Sciences de St-Petersbourg, ser. 6, 5: 12-26 (of

reprint).

WILLEMSE, L.P.M. 1982. A discussion of the Ehrharteae (Gra-

mineae) with special reference to the Malesian taxa former-

ly included in Microlaena. Blumea 28: 181-194.

SPECIMENS EXAMINED

Acocks 2281 7 (1) PRE. Adamson 3603 (2) PRE.

Bolus 4225 (1) BOL, PRE. Bond W662 (1) NBG; 1611 (1) PRE.

Boucher 173 (2) STE; 1838 (2) PRE, STE; 2028 (1) PRE, STE.

Bure hell 7184 ( 1) K.

Bothalia 18,2 (1988)

171

Drige s.n. (1) K, LE (lectotype), P, PRE; s.n. (2) K, LE (lecto-

type), P, PRE, SAM. Du Toit 2027 (1) PRE, STE.

Ecklon s.n. (2) P. Ecklon & Zeyher s.n. (1) LE, SAM; s.n. (2)

LE. Ellis 4695 (1) PRE; 4696 (1) PRE; 5457(1) PRE. Ester-

huysen 7083 (1) BOL; 13657 (1) PRE; 28427 (2) BOL, PRE;

32827a (1) BOL, PRE; 34491 (1) BOL; 35600 (2) BOL, PRE.

Fourcade 2479 (1) BOL, K, PRE, STE; 2852 (1) PRE, STE.

Gill s.n. (1) K.

Haynes 526 (1) K, PRE, STE.

Keet 13-3-1919 (1) PRE. Kensit 14649 (1) BOL.

Ledebour s.n. (2) LE.

MacOwan 2074 (1) SAM; 2126 (1) K.

Paterson 2484 (1) BOL.

Schlechter 3611 (1) PRE; 9809 (1) BOL, K, LE, P, PRE.

Taylor 3221 (1) PRE, STE; 4081 (1) PRE, STE; 4211 (1) JF,

PRE, STE; 5598 (1) PRE; 10256 (1) PRE, STE. Thome Oct.

1926 (1) SAM.

Zeyher 4513 (1) BOL, K, P, PRE, STE; s.n. (1) BOL, LE; s.n.

(2) SAM. Zeyher & Ecklon s.n. (1) P.

SPECIMENS OF THE VILLOSA GROUP OF EHRHARTA

EXAMINED IN BOTHALIA 17,2: 191-194 (1987)

Acocks 14809 (1) PRE; 15185 (1) PRE; 23393 (1) PRE. Adam-

son 1116 (1) PRE; 1240 (2a) BOL; 1360 (2a) PRE; 1362 (2a)

BOL. Andreae 1314 (1) PRE. Archibald 3664 (2b) PRE.

Barker 2718 (2a) NBG. Bohnen 4541 (2a) PRE, STE. Bond 248

(1) NBG; 16 72 (1) NBG; 1685 (1) NBG; 9566 (1) NBG; 9644

(1) NBG. Booysen 2239 (1) NBG. Boucher 662 (2a) STE; 1689

(2b) PRE, STE; 3325 (2a/2b) STE; 3983 (2a) PRE, STE; 4724

(1) PRE, STE. Boucher & Shepherd 5665 (1) PRE. Britten 444

(2b) PRE; 778 (2b) PRE; 1890 (2b) PRE; 1941 (2b) PRE.

Buys 6. 10. 78 (2b) STE.

Cleghom 3122 (2a) PRE, STE. Compton 6262 (1) NBG; 9343

(2a) NBG; 14187 (2a) NBG; 17530 (2b) NBG; 22225(1) NBG,

PRE, STE. Crook 1038 (2a) BOL, NBG, PRE; 2260( 2a) PRE.

Curator, Pretoria 131 (2a) PRE.

Dahlstrand 156 (2b) J, PRE. Dinter 3968 (1) PRE. Dixon 151

(2a) STE. Downing 400 (2a) PRE. Duthie 1642 (1) BOL. Du Toit

1651 (1) PRE, STE; 1668 (1) PRE, STE.

Ecklon Oct. 1838 (1) NBG, SAM. Ecklon & Zeyher 409 (2b)

BOL, NBG, SAM, STE. Ellis 601 (2b) PRE, STE; 708 (1) PRE;

1145 (1) PRE; 1152 (1) PRE; 1284 (2a) PRE; 1651 (l/2a)

PRE; 1686 (2a) PRE; 1700 (1) PRE; 4626 (1) PRE; 4633

(1) PRE; 4635 (1) PRE; 4640 (l/2a) PRE; 4642 (1) PRE;

4648 (1) PRE; 4665 (2a) PRE; 4693 (l/2a) PRE; 5702(1)PRE;

5130 (1) PRE. Emdon 200 (l/2a) PRE, STE. Esterhuysen 3244

(1) PRE; 56 70(1) BOL; 9320 (1) BOL; 22725(1) PRE; 52245

(1) BOL, PRE; 32496 (1) BOL, PRE; 34024 (1) BOL; 34080

(2a) BOL.

Fairale 343 (1) PRE.

Gibbs Russell 5589 (1) PRE; 5591 (1) PRE; 5594 (1) PRE;

5596 (1) PRE; 5607 (1) PRE; 5677 (l/2a) PRE; 5674(1)PRE;

5619 (1) PRE; 5624 (1) PRE; 5628 (1) PRE; 5650 (1) PRE;

5631a (1) PRE; 5638a (1) PRE; 5648 (2a) PRE; 5670 (l/2a)

PRE; 5680 (1) PRE; 5689 (1) PRE. Gluckmann 26/11138 (2a)

J. Grant 4996 (1) PRE.

Hubbard 261 (2a/2b) STE.

Kruger 959 (1) JF, STE; 1681 (1) JF.

Le Maitre 291 (1) JF. Levyns Oct. 1923 (2b) SAM. Liebenberg

4015 (1 /2a) PRE; 4224 (1) PRE; 4229 (2a) STE; 4260 (2a)

STE; 4272 (2a) STE; 4274 (1) STE; 6557 (1) PRE, STE.

Marloth 3046 (2a) PRE, STE; 3712 (2a) PRE; 6024 (2a) PRE.

Merxmueller & Giess 28299 (1) PRE. Moss 3.1.18 (2a) J. M.R.L.

3594 (1) BOL.

Olivier 165 (1) STE.

Pappe Oct. 1838 (1) BOL. Parker 3621 (2a) BOL, NBG. Pater-

son 1097 (2a) BOL. Pearson 5121 (1) BOL, PRE. Phillips 11903

(1) SAM. Pillans 757 9 (1) BOL. Prior April 1903 (2a) SAM.

Schlechter 2098 (1) PRE; 9058 (1) BOL, PRE; 10208 (1) BOL,

PRE. Smith 4969 (2a) PRE. Stocks s.n. (2b) STE. Story 4328

(1) PRE.

Taylor 10167 (2b) PRE, STE. Theron 1108 (2b) PRE. Thoday

Nov. 1921 (2a) BOL. Tyson 607 ( 1) BOL, NBG; 8598 (2b)

PRE, STE; 14870 (2b) BOL; 7 7549 (2b) PRE.

UPE Staff 158 (2b) PRE.

Van Breda 4 (1) PRE. Van Breda & Joubert 2175 (1) STE.

Van der Byl Sept. 1927 (2a) SAM, STE. Van der Merwe 1201

(l/2a) PRE. Van Rensburg 111 (1) PRE, STE; 276 (1) STE.

Walgate 639 (2b) NBG. Walker Dec. 1976 (2a) J. Wolley Dod

3229 (2a) BOL.

Zeyher 167 (1) BOL, PRE, SAM; 4509 (1) PRE; 45096 (2a)

PRE. Zinn 67672 (1) SAM.

Bothalia 18,2: Bothalia 18,2: 173-181 (1988)

The Oxygonum dregeanum complex (Polygonaceae)

G. GERMISHUIZEN*

Keywords: new status, new taxa, Oxygonum dregeanum complex, Polygonaceae, southern Africa, taxonomy

ABSTRACT

The Oxygonum dregeanum Meisn. complex was studied. The following taxa are given a new status: subsp.

swazicum (Burtt Davy) Germishuizen, stat. nov. and subsp. canescens (Sond.) Germishuizen, stat. nov. Four

new varieties of subsp. canescens are described: var. linearifolium Germishuizen, var. nov.; var. dissectum Ger-

mishuizen, var. nov.; var. pilosum Germishuizen, var. nov. and var. lobophyllum Germishuizen, var. nov. Two

new subspecies are described: subsp. streyi Germishuizen, subsp. nov. and subsp. lanceolatum Germishuizen,

subsp. nov.

UITTREKSEL

Die kompleks Oxygonum dregeanum Meisn. is bestudeer. Nuwe status is aan die volgende taksons toegeken:

subsp. swazicum (Burtt Davy) Germishuizen, stat. nov. en subsp. canescens (Sond.) Germishuizen, stat. nov.

Vier nuwe varieteite van subsp. canescens word beskryf: var. linearifolium Germishuizen, var. nov.; var. dissec-

tum Germishuizen, var. nov.; var. pilosum Germishuizen, var. nov. en var. lobophyllum Germishuizen, var. nov.

Twee nuwe subspesies word beskryf: subsp. streyi Germishuizen, subsp. nov. en subsp. lanceolatum Germishui-

zen, subsp. nov.

As part of a revision of the genus Oxygonum Burch,

in southern Africa, to be published in the Flora of south-

ern Africa, the Oxygonum dregeanum complex was

studied. Two subspecies and four varieties are new and

are here described. A list of the changes in rank of the

different taxa of Oxygonum dregeanum are given.

Oxygonum dregeanum Meisn. in Linnaea 14: 487

(1840). Type: neighbourhood of Omsamculo (Umzim-

kulu) (location and collector uncertain).

(a) subsp. dregeanum.

O. dregeanum Meisn.: 487 (1840); C.H. Wr.: 461 (1912);

Tikovsky & Merxm.: 4 (1969); O. dregeanum Meisn. var. dre-

geanum— R.A. Grah.: 166 (1957); Ross: 156 (1972). Type as

above.

O. dregei Meisn.: 38 (1856). Type: Port Natal, Krauss 283

[K, lecto.!, designated by Graham: 166 (1957)].

O. natalense Schltr.: 92 (1907). Type: Natal, Alexandria,

Rudatis 74 (Type not seen).

Found in grassland in sandy soil in Natal and Transkei.

Figure 1.

Vouchers: Huntley 717 (NH, PRE); Johnson 414 (NBG,

STE-U); Strey 8861 (PRE); Wood 628 (GRA, PRE, SAM).

(b) subsp. streyi Germishuizen, subsp. nov., a sub-

specie typica habitu multiramoso, caudice longo cylin-

drico rubro, ocrea subulato-lobata, inflorescentia brevi

differt.

TYPE. — Natal, 3030 (Port Shepstone): Sea Park,

R.G. Strey 11369 (PRE, holo; NH). Figure 2.

Robust, spreading, decumbent perennial herb, much

branched from a long, thick, red vertical rootstock; root-

* Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

MS. received: 1988.01.19.

stock over 200 mm long and 25 mm in diameter. Stems

angular, longitudinally ridged, reddish brown, densely

hairy with white velutinous appressed hairs, rarely glab-

rous; with short internodes. Ocrea membranous, lobed,

reddish brown, densely hairy with white velutinous ap-

pressed hairs, 4—5 mm long; lobes 3,0— 4,5 mm long,

ending in a narrow red tip .Leaves simple, sessile, glabrous

except for white velutinous appressed hairs along the

midrib and base of the leaves; blade elliptic to narrowly

obovate, (8)13—31(32) x (1,5)3—12 mm, acuminate,

ending in a stiff tip, margins crisped. Inflorescence a

short axillary thyrse with fascicles of 1—4 flowers in the

axils of the cup-shaped bracts; axis 40-100 mm long.

Perianth 5-lobed, white, glabrous except for the dense

pilose hairs at the base; the lobes free almost to the base,

5—8 x 2,5 mm, the outer 2 segments keeled at the apex;

pedicels up to 4 mm long, glabrous. Stamens 8, inserted

on the inner perianth; anthers 1 mm long, reddish brown;

filaments 4—5 mm long, with a basal ring of reddish cilia.

FIGURE 1. — Distribution of Oxygonum dregeanum subsp. dre-

geanum.

174

Bothalia 18,2 (1988)

FIGURE 2. — Holotype of Oxygo-

num dregeamm Meisn. subsp.

streyi Germishuizen.

Styles 3, up to 5 mm long, joined for three-quarters of

their length; stigmas capitate. Fruit 9x3 mm, ovoid,

longitudinally ridged, waited between the ridges, veluti-

nous, crowned with the persistent perianth.

Occurs mostly in coastal grassland and palm veld on

sandy soils in Natal. Figure 3.

NATAL. — 2632 (Bella Vista): 40 km from Ndumu on road

to Kosi Bay (-DC), Strey 10305 (NH, PRE). 2732 (Ubombo):

8 km south of Muzi (-AD), Ross <£ Moll 5097 (NH, PRE); IDC

rice project site, 2 km from turnoff to Phelendaba on Mbazwana

road (-BA), Germishuizen 3639 (PRE); just east of Vazi Swamp,

Mazengwenya (-BA), Moll 4741 (PRE); between Lake Vazi

and Sibayi area (-BA), Vahrmeijer 1088 (PRE); 16 km north of

Lake Sibayi on road to Maputo (-BC), Vahrmeijer & Toelken

265 (NH, PRE); Sordwana Bay (-DA), Gerstner 3683 (NH,

PRE); near Bhangazi, Sordwana State Forest ( - D A), MacDevette

1235 (NH, PRE). 2830 (Dundee): Qudeni (-DB), Gerstner

396 7 (NH, PRE). 2832 (Mtubatuba): Richards Bay (-CC),

Venter 4873 (PRE). 2930 (Pietermaritzburg): Inanda (-DB),

Medley Wood 313 (K); Clairmont (-DD), Medley Wood 10497

(NH); Schlechter 2950A (NH). 3030 (Port Shepstone): Sea

Park (-CB), Strey 11369 (NH, PRE); Munster (-CC), Gemmell

sn. (PRE); Uvongo (-CD), Acocks 10907 (PRE); Liebenberg

8102 ( PRE); Mogg 13160 (PRE); Margate, Village of Happiness

(-CD), Coleman 659 (NH); Margate (-CD), Strey 10940, 11023

(NH, PRE); 11069 (NH); St. Michaels on Sea (-CD), Nichol-

son 82 (NH). 3130 (Port Edward): Port Edward (-AA), Nichol-

son 1 712 (PRE); Taylor 5383 (NBG); near Omobaleni (Mgungu)

(-AA), Acocks 13385 (PRE).

This taxon is readily distinguished from the others by

the much branched decumbent habit, the short inter-

nodes, the long, red, cylindrical rootstock and the short

axillary inflorescences.

This subspecies is named in honour of Mr R.G. Strey,

who brought to my attention the fact that this taxon is

different from the typical species.

(c) subsp. swazicum (Burtt Davy) Germishuizen,

stat. nov.

O. zeyheri Sond. var. swazicum Burtt Davy: 167 (1926).

Type: Swaziland, Stewart s.n (K, holo.l).

O. dregeanum Meisn. var. swazicum (Burtt Davy) R. Grah.:

167 (1957).

Bothalia 18,2 (1988)

175

FIGURE 3. — Distribution of Oxygonum dregeanum subsp.

streyi, •; subsp. swazicum, A.

Occurs in eastern and south-eastern Transvaal and

Swaziland (Figure 3). This taxon is distinguished from

the typical subspecies in having deeply linear-lobed leaves

and an unevenly lobed ocrea.

Vouchers: Bolus 12260 (BOL, PRE); Kluge 287 (PRE);

Rogers 11484 (K, NH); Stewarts.ru TM 8880 (GRA, PRE).

(d) subsp. canescens (Sond.) Germishuizen, stat.

nov.

with a small spreading prickle a third of the way from

the base, long and sharp or short and obtuse, obscurely

puberulous.

Five varieties are recognized that can be distinguished

by the lobing of the leaves.

var. canescens

The leaves are deeply divided into 3—9 lobes. Only

the lower lobes are shallowly divided again. Occurs in

open grassland, savanna and thornveld on red sandy soils

in Botswana, Transvaal, northern Orange Free State and

northern Cape Province. Figure 4.

Vouchers: Bayliss 4515 (NBG, PRE); Louw 94 (GRA, PRE);

Morris & Engelbrecht 1070 ( PRE); Pegler991 (BOL, PRE, SAM).

var. linearifolium Germishuizen, var. nov., a varie-

tate typica foliis linearibus integris differt.

TYPE. — Transvaal, Pretoria District, beside freeway

to eastern Transvaal at turnoff to Onbekend, Mauve &

Reid 5326 (PRE, holo.). Figure 5.

A much branched herb with linear leaves, rarely with

1—2 short teeth at the base of lower leaves. Stems and

leaves often dotted with black glands. Occurs in grass-

land, sour bushveld and in disturbed area along road-

sides in the central, southern and south-eastern Trans-

vaal and northern Natal. Figure 6.

O. canescens Sond.: 100(1850); Meisn.: 38 (1856); C.H. Wr.:

462 (1912); Burtt Davy: 167 (1926); Compton: 192 (1976).

O. dregeanum Meisn. var. canescens (Sond.) R.Grah.: 167 (1957).

Type: Transvaal, Aapjesrivier, Zeyher 1451a (W, holo.!).

O. zeyheri Sond.: 100 (1850); Meisn.: 38 (1856); C.H. Wr.:

460 (1912); Burtt Davy: 167 (1926); Compton: 192 (1976).

Type: Transvaal, Magaliesburg, Zeyher 1451b (W, holo.!).

O. calcaratum Meisn.: 38 (1856); Schinz: 870 (1901); C.H.

Wr.: 462 (1912); Burtt Davy: 167 (1926). Type: British Bechu-

analand [northern Cape Province], ‘near the source of Kuruman

river’, Burchett 2459 (K, holo.!).

Annual herb, up to 0,5 m tall, much branched from a

thick vertical perennial woody rootstock. Stems angular,

sulcate, erect, occasionally decumbent, not much swollen

at the nodes, finely to densely white strigose puberulous,

long pilose or rarely glabrous, sometimes dotted with

black glands. Ocreae lobed, up to 5 mm long, finely to

densely strigose puberulous; lobes triangular, ending in

slender red seta, nearly as long as the tube. Leaves (10—)

15— 65(— 70) x (1— )2-20(-26) mm, fleshy, linear,

lanceolate or rhomboid, entire or with 1—2 short teeth

at the base or deeply pinnately or bipinnately lobed; the

lobes linear, further lobed, acute, sometimes with a red

mucro, narrowing and tapering to a short petiole, white

strigose puberulous. Inflorescence a long lax thyrse with

fascicles of up to 3 flowers in the axils of the brown,

ovate cuspidate membranous bracts without seta. Perianth

5-lobed, white, yellow or pinkish; lobes oblong, up to

5 mm long, densely white pilose outside below; pedicels

up to 4 mm long, reddish brown, eglandular, pubescent.

Stamens 8, included; filaments white, up to 5 mm long,

with a ring of red-brown cilia one eighth from the base;

anthers blue, up to 1 mm long. Styles heterostylous, 3,

up to 5 mm long, joined for half their length; ovary pu-

bescent with 3 basal horns. Fruit an obovoid nut, up to

10 x 5 mm, 9-ridged, transversely wrinkled, sometimes

TRANSVAAL.— 2527 (Rustenburg): Jack Scott Private

Nature Reserve (-DC), Wells 2405 (PRE). 2528 (Pretoria):

Great North Road at turnoff to Rust De Winter (-AD), Mauve,

Reid & Smook 5315 (PRE); Kwandebele, Gemsbokfontein

Farm (-BD), Van Hoepen 1680 (PRE); Pretoria (-CA), Rogers

s.ru (PRE); Rooikop (-CA), Smuts & Gittett 2124 (PRE);

Kwaggaspoort (-CA), Verdoom s.n. (PRE); RietvaJlei 221

(— CA), Acocks 11280 (PRE); Koedoespoort (-CB), Werder-

mann & Oberdieck 1244 (PRE); Van der Merwe Station (-CB),

Janse 55 (PRE, SAM); Donkerhoek, 35 km from Pretoria

(—CD); Repton 672 (PRE); 1,8 km S of Rayton on road to

Cullinan (-DA), Davidse 5988 (PRE); Premier Mine (-DA),

Menzies s.ru (PRE); 37 km E of Pretoria (-DC), Dyer & Collett

4680 (PRE); Bronkhorstspruit, Kaalfontein 613 JR Farm (-DC),

Krynauw 985 (PRE); beside freeway to eastern Transvaal af

turnoff to Onbekend (-DC), Mauve & Reid 5326 (PRE). 2529

(Witbank): Groblersdal (-AB), Venter 2971 (PRE); NW side of

Loskopdam (-AC), Reid 686 (PRE); ‘The Hell’, near Loskop-

FIGURE 4. — Distribution of Oxygonum dregeanum subsp. ca-

nescens var. canescens.

176

Bothalia 18,2 (1988)

FIGURE 5. — Holotype of Oxygonum

dregeamim Meisn. subsp. canes-

cens (Sond.) Germishuizen vai.

linearifolium Germishuizen.

dam (-AD), Germishuizen 1392 (PRE); Loskopdam, Twee-

loopfontein (-AD), Theron 1716, 2026 (PRE, PRU); Witbank

(-CC), Rogers 2568 (GRA); between Bronkhorstspruit and

Witbank (— CC), Killick 3839 (PRE). 2530 (Lydenburg): Skur-

weberg, Makobulaan (-BA), Onderstall 1201 (PRE). 2628

(Johannesburg): Modderfontein (-AA), Prosser s.n. (NBG,

PRE); 43 km SE of Pretoria, 1 mile (1,6 km) E of Bapsfontein

(-BB), Codd 2575 (PRE). 2629 (Bethal): Breyten (-BD), Steyn

999 (NBG).

NATAL. — 2730 (Vryheid): Mooihoek (—AC), Devenish

938 (PRE); 11 km from Vryheid on road to Louwsburg (— DD),

Germishuizen 2260 (PRE).

var. dissectum Germishuizen , var. nov., a varietate

typica lobis profundioribus, pluribus, et apicibus (lobo-

rum) acutis differt.

TYPE. — Transvaal, Pietersburg, Feb. 1904, H. Bolus

s.n. (BOL, holo.; GRA). Figure 7.

The leaves of this plant are deeply divided and the FIGURE 6. — Distribution of Oxygonum dregeanum subsp. ca-

lobes in turn are again divided; each lobe ending in a nescens var. linearifolium.

Bothalia 18,2 (1988)

111

long slender tip. Occurs in grassland in sandy soils in the

northern Transvaal, especially in the Pietersburg area.

Figure 8.

TRANSVAAL. — 2329 (Pietersburg): Blinkwater (-AA),

Bremekamp & Schweickerdt 188 (PRE); Pietersburg (-CD),

Bolus s.ru (BOL, GRA); Pietersburg Nature Reserve (-CD),

Bredenkamp & Van Vuuren 123 (PRE); Marabastad (-CD),

Nelson 118 (PRE). 2428 (Nylstroom): Naboomspruit, Mosdene

(-DA), Galpin M305 (PRE, SAM). 2429 (Zebediela): Percy

Fyfe Nature Reserve (-AA), Huntley 1146, 1550 (PRE); Luns-

khp, about 32 km NNE of Potgietersrust (-AA), Maguire 1370

(NBG).

var. pilosum Germishuizen, var. nov., a varietate

typica lobis inferioribus foliorum longioribus et plantis

pilosis differt.

TYPE.— Transvaal, Twentyfour Rivers, F.A. Rogers

23637 (PRE, holo.). Figure 9.

This taxon differs from the typical variety by the

lower lobes of the leaves being over 1 5 mm in length,

and the whole plant covered with long white pilose hairs.

Occurs only in the north-western Transvaal, especially in

the Waterberg area in savanna on sandy soils. Figure 8.

TRANSVAAL. — 2428 (Nylstroom): Alma, 56 km NW of

Warmbaths (-AC), Bayliss 2007 (NBG); Van Rensburg Farm,

43 km along Nylstroom-Vaalwater road (-AC), Leeuwenberg

10900 (PRE); Waterberg, Vaalwater, Farm Nooitgedacht (-AC),

Westfall 2279 (PRE); Waterberg, Zandrivierspoort near Alma on

Farm Witpoort (-AC), Van Wyk 26 79 (PRE, PRU); Twenty-

four Rivers (-AD), Rogers 23637 (PRE); Geelhoutkop (-AD),

Breyers.ru (PRE).

var. lobophyllum Germishuizen, var. nov., a varie-

tate typica lobis foliis latioribus et lobis ocreae triangu-

laribus differt.

TYPE. — Transvaal, 18 km ENE of Pretoriuskop,

Acocks 16643 (PRE, holo.). Figure 10.

The leaves are shallowly and deeply lobed but not di-

vided up to the midrib, rarely entire; lobes usually wider

than 2 mm. Occurs in the northern and eastern Transvaal

and Swaziland in woodland, bushveld and grassland in

sandy loam or granitic soils. Figure 8.

FIGURE 7. — Holotype of Oxygonum dre-

geanum Meisn. subsp. canescens (Sond.)

Germishuizen var. dissectum Germis-

huizen.

Austro Africanae.

no iom.

.OJU^OiUcuv lUzjlvru-, Sc-tvL

, £-r H-t

Oxygonum dregeanum

canescens (Sond.) <

var. dissectum Ger:

178

FIGURE 8. — Distribution of Oxygonum dregeanum subsp. ca-

nescens vai.dissectum, A; vai.pilosum, 4; vai.lobophyllum.

Bothalia 18,2 (1988)

TRANSVAAL. — 2230 (Messina): Venda, Ha-Lambani (-DD),

Mugwedi 1665 (PRE). 2329 (Petersburg): 13 km from Bandoliers-

kop on Petersburg road (-BD), Story 1833 (PRE). 2330 (Tza-

neen): Hans Merensky Nature Reserve (-DA), Oates 88, 128

(PRE). 2431 (Acomhoek): Farm Kempiana 90 KU, Timbavati

Private Nature Reserve (-AD), Zambatis 1470 (PRE); Pilgrims

Rest District, Manyeleti Game Reserve (-DA), Bredenkamp

1262, 1382 (PRE). 2530 (Lydenburg): Nelspruit (-BD), Breyer

s.n. (PRE); Nelspruit Research Station (-BD), Liebenberg 2152

(PRE). 2531 (Komatipoort): Kruger National Park, 18 km ENE

of Pretoriuskop (-AB), Acocks 16643 (PRE); near Pretoriuskop

(- AB), Codd & De Winter 4933 (PRE); 10 km N of Pretorius-

kop on road to Skukuza (-AB), Codd & De Winter 4993, 4994

(PRE); Nohpe (-AB), Van der Schijff 1541 (PRE); Kaapmui-

den (— CB), Rogers s.ru (PRE).

SWAZILAND. —2631 (Mbabane): Malinda Hill (-AC),

Compton 28433, 29775 (NBG, PRE); Stegi District, Sicusha

(-BC), Compton 32472 (NBG, PRE).

(e) subsp. lanceolatum Germishuizen , subsp. nov.

a subspecie typica foliis angustioribus et tepalis vinoso-

rubris differt.

TYPE. — Transvaal, Lekgalameetse Nature Reserve,

The Downs, Makwens, M. Stalmans 192 (PRE, holo.).

Figure 1 1 .

FIGURE 9. — Holotype of Oxygonum dre-

geanum Meisn. subsp. canescens (Sond.)

Germishuizen var. pilosum Germishui-

zen.

Bothalia 18,2 (1988)

179

FIGURE 10. — Holotype of Oxy-

gonum dregeanum Meisn.

subsp. canescens (Sond.)Ger-

mishuizen var. lobophyllum

Germishuizen.

7 'K / ttt / ,

J, }1 ff C(c.ocs * /&. iS

c 'Zoo'

Z3///53

liTwtn

Robust herb, arising from a thick vertical perennial

woody rootstock, with many erect annual stems up to

0,3 m long. Stems much branched, angular, sulcate, fine-

ly and densely strigose puberulous, rarely glabrous,

sometimes reddish tinged. Ocrea up to 8 mm long, lobed,

membranous, brown, sometimes with a reddish tinge,

white strigose puberulous, rarely glabrous; lobes up to

4 mm long, ending in a narrow red tip. Leaves (15—)

20— 33(— 40) x (1 — )1 ,5 — 3,5(— 8) mm, lanceolate to

narrowly ovate, rarely lobed towards base, glabrous ex-

cept for a few white appressed hairs along the midrib at

base and along margins, rarely white strigose puberulous,

acute, all ending in a stiff point, tapering to narrow

petiole, mostly with dark red margins especially near

apex. Inflorescence an axillary or terminal thyrse with

up to 3 flowers in brown, cuspidate, cup-shaped bracts;

axis 50—200 mm long. Perianth 5-lobed, white, deep

pink or wine-red tinged along centre on outside of outer

3 lobes, densely pilose at base becoming glabrous towards

the apex; lobes free almost to base, up to 7 mm long,

outer 3 lobes keeled at apex; pedicels strigose, up to 4

mm long. Stamens 8, inserted on inner perianth lobes;

filaments of different lengths, up to 5 mm long; anthers

up to 1,5 mm long, reddish brown, with a ring of red-

dish brown cilia at base. Styles 3, heterostylous, 2,5—

6,0 mm long, joined for less than half their length. Fruit

an ovoid nut, up to 12 mm long, slightly 3-angled, usual-

ly without 3 blunt teeth at base, 9-ridged, transversely

wrinkled, glabrous.

Found in open grassland and on montane rocky grass-

land, or rarely in woodland in Transvaal, Swaziland and

Natal. Figure 12.

TRANSVAAL. — 2430 (Pilgrims Rest): L^kgalameetse Nature

Reserve, Makwens (-AA), Stalmans 192 (PRE); Haffenden

Heights (-AA), Stalmans 174 (PRE); road to Makwens (-AA),

Stalmans 634 (PRE); The Downs (-AA), Junod 4253 (PRE),

Stalmans 1452 (PRE); on top of Orrie Baragwanath Pass (-AA),

Reid 1159 (PRE); Ohrigstad Dam Nature Reserve (-DC),

Jacobsen 1769 (PRE); Mount Sheba Nature Reserve, N slope

of Mount Sheba (-DD), De Winter 9361 (PRE); Graskop

(-DD), Galpin 14569 (PRE); Graskop, Stanley Bush Kop

(-DD), P. & G. Raal 1035 (PRE); Pilgrims Rest (-DD), Rogers

14314 (PRE). 2530 (Lydenburg): Boschhoek, western Steen-

180

Bothalia 18,2 (1988)

Oxygonum dregeanum Melsn. subsp.

lanceolatum Germishuizen

"i 1988

$ metOW

HERBARIUM PRETORIA

f,5G053

FIGURE 11. — Holotype of Oxygo

mm dregeamm Meisn. subsp.

lanceolatum Germishuizen.

FIGURE 12. — Distribution of Oxygonum dregeanum subsp.

lanceolatum.

kampsberg (-AA), Young A407 (PRE); Hartebeestvlakte (-B A),

Mohle 151 (PRE); Mauchsberg, Sabie (-BB), Smuts & Gillett

2306 (PRE, STE-U); Wonderkloof Nature Reserve (-BC), Elan-

Puttick 128 (PRE); Belfast (-CA), Jenkins s.n. (PRE). 2531

(Komatipoort): Barberton (-CC), Pott 5469 (PRE); Thorn-

croft s.n. (PRE); Saddleback Mountain (-CQ, Galpin 615

(NH, PRE, SAM); Osterbeek Farm, 5 km S of Barberton (-CC),

De Sousa 626 (PRE); Swaziland border, Daylight Farm, Bearded

Man Mountain {-CD),Buitendag 756 (NBG, PRE). 2629 (Bethal):

Ermelo (— DB), Riley 12 (PRE).

SWAZILAND. — 2531 (Komatipoort): Havelock Mine, Piggs

Peak (-CC), Miller 2988 (PRE). 2631 (Mbabane): Malolotja

Nature Reserve, Nkhaba (-AA), Heath 470 (PRE); Black Mbu-

luzi Valley (-AA), Dlamini s.n. (NBG, PRE); Ukutula (-AC),

Compton 25256 (NBG, PRE); Compton 24501 (NBG).

NATAL. — 2731 (Louwsburg): 15 km from Ngome Forest

Station on road to Vryheid (-CD), Germishuizen 2133 (PRE);

12 km from Ngome Forest Station on road to Vryheid (-CD),

Buthelezi 115 (NH, PRE). 2829 (Harrismith): Klip River, Stryd-

hoek, Tintwa Mountains (— CB), Doidge s.n. (PRE). 2930 (Pieter-

maritzburg): Tweedie (-AC), Curator Pretoria 3462 (PRE);

Greytown (-BA), Wylie s.n. (PRE); Pietermaritzburg, Table

Mountain (-CB), Killick 732 (NBG).

Bothalia 18,2 (1988)

ACKNOWLEDGEMENTS

My thanks go to the following: the curators and staff

of BOL, GRA, K, NH, NBG, PRE, SAM, STE-U and W

for the loan of their material; Dr H.F. Glen for translating

the diagnoses into Latin; and Mrs A.J. Romano wski for

the photographs.

REFERENCES

BURTT DAVY, J. 1926. A manual of the flowering plants and

ferns of the Transvaal with Swaziland, South Africa 1:

167. Longmans, Green, London.

COMPTON, R.H. 1976. Polygonaceae. Flora of Swaziland.

Journal of South African Botany, Supplement Vol. 11:

192.

GRAHAM, R.A. 1957. A revision of Oxygonum. Kew Bulletin

1: 166-167.

181

MEISNER, C.F. 1840. Polygoneae. In D.F.L. Schlechtendal,

Linnaea 14: 487.

MEISNER, C.F. 1856. Polygonaceae. In A.P. De Candolle,

Prodromus systematis naturalis regni vegetabilis 14: 38.

Paris.

ROSS, J.H. 1972. Flora of NataL Memoirs of the Botanical

Survey of South Africa No. 39: 156. Government Printer,

Pretoria.

SCHINZ, H. 1901. Polygonaceae. Bulletin de I’Herbier Boissier

ser. 2,1: 870.

SCHLECHTER, R. 1907. Beitrage zur Kenntnis der Flora von

Natal. In H.G.A. Engler, Botanische Jahrbucher 40: 92.

Leipzig.

SONDER, O.W. 1850. In D.F.L. Schlechtendal, Linnaea 23 : 100.

Halle, Berlin.

TIKOVSKY, H. & MERXMULLER, H. 1969. Polygonaceae. In

H. Merxmuller, Prodromus einer Flora von Sudwest-afrika

23: 4. Cramer, Lehre.

WRIGHT, C.H. 1912. Polygonaceae. In W.T. Thiselton-Dyer,

Flora capensis 5,1: 461-462. Reeve, London.

Bothalia 18,2: 183-188 (1988)

Notes on African plants

VARIOUS AUTHORS

BRASSICACEAE

HELIOPHILA CORNELLSBER GIA , A NEW SPECIES FROM THE RICHTERSVELD

Heliophila comellsbergia B.J. Pienaar & A. Nicholas

sp. nov. H. patenti Oliv. et H. diffusae (Thunb.) DC. var.

diffusae et vai.flaccae (Sond.) Marais affrnis; a H. patenti

floribus multo maioribus et fructu globoso, inflato dif-

fert; a H. diffusae et varietatibus foliis non divisis differt.

Herba annua. Caulis unum ad aliquot. Folia simplices,

integra, estipulata, anguste-lanceolata ad elliptica, 10-

40 x 2—8 mm. Flores albi. Petala 2,9— 4,5 x 1,4— 2,5

(—2,8) mm. Fructus globosus, 3,0— 4, 0(— 4,5) x 3,0—

3,8(— 4,0) mm, valvis inflatis.

TYPE.— 2817 (Vioolsdrif): (-CA) Richtersveld,

Cornellsberg in Stinkfontein Mountains, southern slopes

to a neck south of the top, 1977.09.06, Oliver, Tolken &

Venter 715 (PRE, holo.; K, MO).

Soft, patently hairy, annual herb, (110— )1 55— 290

mm high. Stems one to several, erect, lax, patently pu-

bescent. Leaves simple, alternate, entire, exstipulate,

narrow-lanceolate to obtusely elliptic, marginally pu-

bescent, midrib ventrally visible, 10—40 x 2—8 mm,

larger leaves near the stem base, both stem and leaves

often tinted purple. Racemes few to many-flowered, lax,

axis elongating in fruit. Pedicels in flower 3—11 mm

long, in fruit 6 — 10( — 12) mm long, initially ascending,

recurved (almost drooping) when mature, patently pu-

bescent. Flowers white, drying cream. Sepals (2,0—)

2,1— 2,5 x 1,0— 1,4 mm, membranous, dark cream in

colour; outer 2 cucullate, inner 2 saccate, margin broad-

ly transparent, tinted purple apically. Petals 2,9— 4,5 x

1,4— 2,5(-2,8) mm, obovate above, claw with lateral

papillate appendage. Filaments 1,6— 1,9 mm and 1,2—

1,7 mm long, the 2 shorter filaments basally papillate.

Anthers (0,5-)0,6-0,7(-0,8) mm long. Ovary globose,

0,4— 0,6 mm in diameter, 2 ovules, style as long as ovary;

stigma capitate, papillate. Fruits globose, 3,0— 4, 0(— 4,5)

x 3,0— 3, 8(— 4,0) mm, puberulent, valves inflated, faint-

ly 1 -nerved, tinged purple when mature; style persistent,

filiform, 2,0— 3,0 mm long. Seeds 1 or 2, 0,8— 1,0 mm in

diameter, narrowly winged. Figures 1 & 2.

CAPE. — 2817 (Vioolsdrif): Rosyntjiesberg, (-AC), Oliver,

Tolken & Venter 302 (PRE); Cornellsberg (-CA), Oliver, Tolken

& Venter 715 (K, MO, PRE).

Heliophila comellsbergia is endemic to mountainous

areas of the Richtersveld. Found in stoney places, in

well drained clayey loam soils, it occurs on the southern

slopes of mountains where it may grow in open or shaded

areas. Oliver, Tolken & Venter record it as locally com-

mon. The Richtersveld is a mountainous area that bor-

ders on the Namib Desert. For much of the year it is

a harsh, extremely dry area in which only succulent

plants and plants with underground storage organs sur-

vive. Good rains in August-September transform parts

of the Richtersveld into a carpet of flowers, as in Nama-

qualand. It is at this period that one finds an abundance

of annuals, such as Heliophila comellsbergia.

The specific epithet comellsbergia is derived from the

isolated locality in which this new Heliophila species was

collected. Cornellsberg is a peak in the Stinkfontein

Mountain range. Oliver, Tolken & Venter collected the

type specimen on the southern slopes, towards the neck

of this peak.

Y h

FIGURE 1. — Holotype {Oliver, Tolken & Venter 715) of Helio-

phila comellsbergia housed at PRE (Photograph by A. Ro-

manowski).

184

FIGURE 2. — 1, petal with claw

and basal appendage, X 1 3 ;

2, stamens, 2a, longer sta-

men without basal papillae,

X 29; 2b, shorter stamen

with basal papillae, X 30;

3, mature globose fruit, X

3. All drawings based on

Oliver, Tolken & Venter

302 (PRE). Drawings by

G. Condy.

Although the material of this new Heliophila species

is limited, the authors find it desirable to name this taxon

for two reasons; a, the remoteness of the area in which

this species occurs, making it unlikely that it will be col-

lected again in the near future; b, the distinctive, unique

morphology of H. comellsbergia.

Bothalia 18,2 (1988)

Heliophila comellsbergia most closely resembles H.

patens Oliv., another Heliophila species of restricted dis-

tribution. H. patens has so far only been recorded from

the Piketberg and a collecting site somewhere between

Knechtsvlakte (this name is probably an orthographic

error and should be Knersvlakte) and Sandveld near Van-

rhynsdorp (Figure 3). However, H. comellsbergia differs

from H. patens in a number of structurally important

and taxonomically significant ways. Of particular note

are the larger flowers, recurved, inflated, globose, 1 or 2-

seeded fruits and pubescent stems, leaves, pedicels and

fruits of H. comellsbergia; H. patens has obovate to

subcircular, laterally flattened fruits with at least two

seeds (Table 1; Figure 4). Although H. comellsbergia

appears to be allied to H. patens, the relationship between

these two species is probably distant. Using Marais’s

(1970) key to Heliophila in the Flora of southern Africa,

H. comellsbergia keys out to H. diffusa (Thunb.) DC.,

FIGURE 3. — Distribution of Heliophila comellsbergia, •; H.

patens, <>; H. diffusa var. diffusa, ■; and H. diffusa var.

flacca, A.

TABLE 1 . — Comparison of selected characters in Heliophila comellsbergia, H. patens, H. diffusa var. diffusa and H. diffusa var. flacca

important diagnostic characters.

Bothalia 18,2 (1988)

185

FIGURE 4. — Scatter diagram (leaf width X fruit length) illus-

trating the distinctness of Heliophila cornellsbergia, <>;

H. patens, o; H. diffusa var. diffusa, ■; and H. diffusa

var. flacca, A . Solid symbols = pinnately compound leaves,

unfilled symbols = simple undivided leaves.

which has two varieties, viz. var. diffusa and var. flacca

(Sond.) Marais. This species differs from H. cornellsbergia

in having pinnately compound leaves, 1— 8-seeded, flat

fruits and glabrous vegetative parts. Although allied to

H. diffusa, the relationship is not considered close.

In his southern African flora treatment, Marais (1970)

places H. patens, H. diffusa var. diffusa and var. flacca

following one another, and there is little doubt that H.

cornellsbergia should be placed with this complex. It is

probable that a long period of isolation on the mountains

of the Richtersveld could explain why it differs fairly

markedly from its nearest relatives, which occur some

distance to the south (Figure 3).

SPECIMENS EXAMINED

Heliophila cornellsbergia B.J. Pienaar & A. Nicholas

CAPE. — 2817 (Vioolsdrif): Rosyntjiesberg (-AC), Oliver,

Tolken & Venter 302 (PRE); Comellsberg (-CA), Oliver, Tol-

ken & Venter 715 (PRE, holo. ; K, MO, iso.).

Heliophila patens Oliv.

CAPE. — 3218 (Clanwilliam): Piketberg (-DD), Bolus 7530*

(BOL, NBG, PRE, SAM, isotypes), Schlechter 3482 (PRE),

5183 (BOL). Without precise locality: between Knechtsvlakte

& Sandveld (possibly 3118BC?), Leipoldt 3963* (BOL, PRE).

Heliophila diffusa (Thunb.) DC. var. diffusa

CAPE. — 3218 (Clanwilliam): Elandskloof (-BD), Adamson

s.n. (BOL 27121), Lewis s.n. (BOL 27120). 3318 (Cape Town):

Kapokberg (- AD), Bolus 12599* (PRE); Lions’ Head & Rump

(—CD), Lilians 3924* (PRE), Ecklon & Zeyher 71* (SAM);

Devils Peak (—CD), Ecklon & Zeyher 70* (SAM); Hercules

Pillar (— DB), Compton 13673* (NBG); Tigerberg (-DC),

Esterhuysen 17509 (PRE); Groot Drakenstein Mts, Duiwels-

kloof (-DD), Esterhuysen 14046* (BOL); Seven Sisters Mtn,

Paarl (-DD), Esterhuysen 23928 (BOL). 3319 (Worcester):

Kleinvlei Bridge (-AB), Acocks 19884* (PRE); Elandskloof

(-AC), Compton 16179 (NBG); Steendal, Tulbagh (-AC),

Zeyher s.n. (SAM 14059)*; Theronsberg (-BC), Compton

11788* (NBG). 3418 (Simonstown): Constantiaberg (-AB)

Compton 13916 (NBG). Precise locality unknown: Ecklon &

Zeyher s.n. (PRE 23076); Braakfontein (?), Schlechter 3840

(PRE).

Heliophila diffusa (Thunb.) DC. var. flacca

CAPE. — 3318 (Simonstown): Helderberg (-BB), Parker

4247* (NBG); Jonkershoek (-DD), Compton 15339* (NBG),

Lewis 3132* (SAM). 3419 (Caledon): Grabouw (— AA), Stokoe

s.n* (SAM 64258); Caledon (-AB), Ecklon & Zeyher 69* (K

142686, holo.; SAM, iso.); Caledon near hotsprings (-AB),

Zeyher s.n. (K 142686), Zwarteberg, Zeyher 1894 (PRE).

ACKNOWLEDGEMENTS

The authors would like to thank the Director and

staff (particularly Dr H. Glen, G. Condy, A. Romanowski,

B. Schrire and J. van Rooy) of the Botanical Research

Institute. The Directors and staff of BOL, K, NBG, and

SAM are thanked for the loan of specimens. The referees

are also thanked for their valuable contribution.

REFERENCE

MARAIS, W. 1970. Cruciferae. In Flora of southern Africa 13:

1 — 118. Government Printer, Pretoria.

B.J. PIENAAR and A. NICHOLAS

* Specimen cited in Flora of southern Africa by Marais (1970).

MS. received: 1988.03.28.

ERICACEAE

COILOSTIGMA ZEYHER1ANUM — A CORRECTION

In ‘Studies in the Ericoideae (Ericaceae). V. The genus

Coilostigmd in Bothalia 17,2: 167, the variety tenui-

folium was published as a new combination without the

citation of the basionym which was inadvertently left

out in the editorial process. This is now rectified.

Coilostigma zeyherianum Klotzsch var. tenuifo-

lium (Klotzsch) E.G.H. Oliver, comb, et stat. nov.

Coilostigma tenuifolium Klotzsch in Linnaea 12: 234 (1838).

Types: In planitie inter ‘Krakakamma’ et montes ‘Vanstadens-

riviersberge’, Ecklon & Zeyher s.n. (Bt, E!, S!); idem as 294

(G!, GOET!, LD!, M!, MEL!, MO!, S!, W!, UPS!, Z!); in sylvis

‘Olifantshoek’ prope flumen ‘Bosjesmansrivier’, Ecklon & Zeyher

s.n. (Bt, BOL!). Lectotype (chosen here): Ecklon & Zeyher s.n.

(S).

E.G.H. OLIVER

186

Bothalia 18,2 (1988)

GERANIACEAE

THE CORRECT AUTHOR CITATION FOR PELARGONIUM SECTION OTIDIA

In the most recent taxonomic treatment of the genus

Pelargonium, Knuth (1912: 369) cited the authority of

the section Otidia as (Lindl.) Harv. (1860), based on the

genus Otidia Lindl. ex Sweet. This is not correct.

The name Otidia was first published, in generic rank,

by Sweet (1820: viii), without any mention of Lindley.

Later in the same work, under t. 98 Sweet stated: ‘This

plant [Otidia carnosa] is proposed by Mr Lindley to form

a distinct genus, which we have adopted ...’. In my opinion

Sweet is the undoubted author of the genus, and ‘Lindl.

ex’ should be omitted from the citation as Lindley was

not mentioned in connection with the original publica-

tion of the name.

Furthermore, Harvey (1860: 278) was not the first

author to place Otidia in sectional rank, being preceded

by G. Don (1831: 729). Don did not cite the generic

name Otidia as basionym, but by citing ‘Lindl. in Sweet,

ger. p. 8 no. 98’ he provided an adequate reference to

the validly published basionym. Although this reference

is not entirely direct, it is not contrary to I.C.B.N. Art.

33.2 which only concerns publications later than 1 Jan.

1953. (A change in rank is considered to be a different

combination and there is not a separate clause to cover

changes in ranking).

The correct author citation therefore is Pelargonium

sect. Otidia (Sweet) G. Don.

REFERENCES

DON, G. 1831. A general system of gardening and botany, Vol.

1. Rivington, London.

HARVEY, W.H. 1860. Pelargonium. In W.H. Harvey & O. Son-

der, Flora capensis 1: 259-308 (278). Hodges & Smith,

Dublin.

KNUTH, R. 1912. Pelargonium. In A. Engler, Das Pflanzenreich

4, 129, 53: 316-545 (369). Engelmann, Berlin.

SWEET, R. 1820. Geraniaceae, Vol. 1. Ridgway, London.

P. VORSTER*

* Botany Department, University of Stellenbosch, Stellen-

bosch 7600, South Africa.

MS. received: 1987.08.22.

ORCHIDACEAE

ADDITIONS TO THE SYNONYMY OF EULOPHIA SCHWEINFURTHII

For some time doubt has existed as to the precise

nature of Eulophia chrysops Summerh., particularly

with regard to its alliance with a species recognized as

occurring in southern Africa, E. schweinfurthii Kraenzl.

(Kraenzlin 1893; Summerhayes 1958; Hall 1965).

Both these concepts have wide distributions in tropi-

cal Africa, extending marginally into the Transvaal (E.

schweinfurthii) and in recent records, into Botswana

(E. chrysops ). Summerhayes (1958) proposed E. chrysops

as a new name for Lissochilus aurantiacus Reichb. f.

(1865), E. aurantiaca having been used previously for

another species, related to E. welwitschii (Reichb. f.)

Rolfe. Summerhayes also reduced two tropical species,

L. johnstonii Rolfe and L. holubii Rolfe to the synonymy

of E. chrysops.

In discussing E. chrysops, Summerhayes (1958) noted

at the time that more data and material were needed to

elucidate its relationships. The specimens that he had

seen suggested that the flowers of E. chrysops were ap-

preciably larger (petals 13—16 mm long) than those of

E. schweinfurthii and that they have less acute sepals

with more narrow bases. Subsequent examination of

material has shown that these distinctions do not hold,

either for the sepal shapes or for the flower sizes. Petal

length varies from 8— 14(— 16) mm in E. schweinfurthii,

covering the range given by Summerhayes fori?. chrysops.

Some of the variation in flower size may be related to

the habitats which vary from arid mopane woodland to

wet bog verges (Williamson 1977) and to dry bushveld

and grassland (Hall 1965). The overall evidence of con-

tinuity shows convincingly that the two concepts should

be merged.

The name of the joint concept should be E. schwein-

furthii Kraenzl. (1893) which, being earlier, replaces

Summerhayes’ E. chrysops (1958) as the nomen novum

for L. aurantiacus Reichb. f. (1865). Further accretions

to this concept are likely to become evident in studies of

material of other tropical allies but none appear to have

been described earlier than E. schweinfurthii.

REFERENCES

HALL, A.V. 1965. Studies of the South African species of

Eulophia. Journal of South African Botany Supplemen-

tary Vol. 5: 1-248.

KRAENZLIN, F. 1893. Orchidaceae africanae. Botanische Jahr-

biicher 17: 54.

REICHENBACH, G.L. 1865. Dr. Welwitsch’s Orchideen aus

Angola. Flora 48: 177-191.

SUMMERHAYES, V.S. 1958. African orchids, 25. Kew Bulle-

tin 1958: 57-87.

WILLIAMSON, G. 1977. The orchids of South Central Africa,

pp. 237. Dent, London.

A.V. HALL*

* Bolus Herbarium, University of Cape Town, Rondebosch 7700.

MS. received: April 1988.

Bothalia 18,2 (1988)

POLYGONACEAE

REINSTATEMENT OF OXYGONUM ACETOSELLA WELW.

187

R.A. Graham in his revision of Oxygonum in Kew

Bulletin 1957: 165 (1957), places Oxygonum acetosella

Welw. as a synonym of Oxygonum alatum Burch, because

they are similar in their winged fruits and lobed leaves.

On closer investigation it is found that O. acetosella dif-

fers from O. alatum by the total absence of cup-shaped

and elongated scales scattered over the entire surface of

the plant, the shallowly lobed leaves and the entire ocreae,

which are always fringed in the latter. Figure 5 repre-

sents a scatter diagram, using internode length along one

axis and width of leaves along the other axis, which

clearly separates the two taxa.

3a

O. acetosella Welw. in Transactions of the Linnean

Society of London 27: 60 (1869); Hiern: 903 (1900);

Bak. & C.H. Wr.: 99 (1909); Gossweiler: 395 (1953).

Type: Angola, Mossamedes, Welwitsch 1757 (BM iso.; K;

PRE!). Figure 6.

A glabrous, somewhat fleshy, glaucous, decumbent or

prostrate annual herb, much branched from the base.

FIGURE 5. — Scatter diagram illustrating the difference between

Oxygonum acetosella, •, and O. alatum, 6.

FIGURE 6. — Type of Oxygonum

acetosella Welw.

r

Oxygonum acetosella Welw.

FACULDADE° DC "-r An9°'- "' 17571

Oxygonum acetosella Welw.

Estafao Angola: Mossamedes

Habitat

, Alt. .. Dat. VI-V1I-1859

Col. F.. Welwitsch N* 1757

Del. Obs.

188

Bothalia 18,2 (1988)

Stems semicylindrical, glabrous, up to 200 mm long.

Ocrea truncate, up to 8 mm long, quite entire or minute-

ly erose-denticulate, with a green leaf-like limb up to 3

mm long. Leaves broadly lanceolate to ovate, simple or

shallowly 2-3-lobed, (30-)39-56(-60) x (9-)12-22

(—27) mm, acute at the apex, narrowed into the winged

petiole, fleshy, glabrous. Inflorescence a small thyrse

with fascicles of up to 3 flowers in the axils of ovate

membranous bracts. Flowers bisexual. Perianth segments

5, obovate, 5—6 mm long, white. Stamens 8; anthers

pale blue, up to 1 mm long; filaments 2, 5-4,0 mm long

with a ring of cilia one-eighth from base. Styles 3, joined

for almost half their length, up to 5 mm long. Fruit ob-

long, up to 10 mm long, with 3 narrow wings.

Found on grassy flats and sandy hills in northern

South West Africa/Namibia. Figure 7.

Vouchers: Giess 7954 (PRE, WIND); Moss & Jacobsen

K299 (PRE).

REFERENCES

BAKER, J.G. & WRIGHT, C.H. 1909. Polygonaceae. In W.T.

Thiselton-Dyei, Flora of tropical Africa 6,1: 99. Reeve,

Kent.

GOSSWEILER, J. 1953. Nomes indigenas de plantas de Angola'.

395. Luanda.

GRAHAM, R.A. 1957. A revision of Oxygonum. Kew Bulletin

1: 165.

FIGURE 7. — Distribution of Oxygonum acetosella Welw. in

South West Africa/Namibia.

HIERN, W.P. 1900. Polygonaceae. Catalogue of the African

plants collected by Dr Friedrich Welwitsch in 1853-1861 :

903. London.

WELWITSCH, F. 1869. Polygonaceae. Transactions of the Lin-

nean Society of London 27 : 60. London.

G. GERMISHUIZEN

MS. received: Feb. 1988.

Bothalia 18,2: 189-194 (1988)

Fusarium tricinctum (Fungi : Hyphomycetes) in South Africa —

morphology and pathogenicity*

SANDRA C. LAMPRECHT,** W.F.O. MARASAS,1- P.S. VAN WYK,* and P.S. KNOX-DAVIES0

Keywords.' Fusarium tricinctum, Medicago truncatula, morphology, pathogenicity, soil, South Africa, taxonomy, Triticum aestivum

ABSTRACT

The occurrence, cultural characteristics, morphology and pathogenicity of seven isolates of Fusarium tricinc-

tum (Corda) Saccardo, are described. Cultures were isolated from plant debris in soils under natural grassland in

the Golden Gate National Park, Orange Free State, and under either wheat or annual Medicago spp. at Malmesbury,

Cape Province. One isolate was obtained from roots of a Medicago sp. at Caledon, Cape Province. The isolates

were identical to each other and to two foreign reference cultures of F. tricinctum and all produced napiform

to citriform microconidia on monophialides. All the isolates were weak to intermediately aggressive pathogens

of Medicago truncatula Gaertn. cv. Jemalong and Triticum aestivum L. cv. Palmiet and caused pre- and post-

emergence damping-off of seedlings as well as discolouration and necrosis of root and crown tissues. This is the

first report of the pathogenicity of F. tricinctum to M. truncatula.

UITTREKSEL

Die voorkoms, kultuureienskappe, morfologie en patogeniteit van sewe isolate van Fusarium tricinctum

(Corda) Saccardo, word beskryf. Kulture is uit plantreste in gronde onder natuurlike grasveld in die Golden

Gate Nasionale Park, Oranje-Vrystaat, en onder of koring of eenjarige Medicago spp. in Malmesbury, Kaap-

provinsie, gei'soleer. Een isolaat is verkry vanuit wortels van ’n eenjarige Medicago sp. te Caledon, Kaapprovin-

sie. Die isolate was identies aan meekaar en aan twee buitelandse verwysingskulture van F. tricinctum en almal

het napiforme tot sitriforme mikrokonidia op monofialides geproduseer. A1 die isolate was swak tot middelmatig

aggressiewe patogene van Medicago truncatula Gaertn. cv. Jemalong en Triticum aestivum L. cv. Palmiet en het

voor- en na-opkomsafsterwing van saailinge sowel as verkleuring en nekrose van wortel- en kroonweefsel veroor-

saak. Hierdie is die eerste aanmelding van die patogeniteit van F. tricinctum teenoor M. truncatula.

INTRODUCTION

Four species are currently accepted in the section

Sporotrichiella of the genus Fusarium Link, i.e. F. poae

(Peck) Wollenw., F. tricinctum (Corda) Sacc., F. sporo-

trichioides Sherb., and F. chlamydosporum Wollenw. &

Reinking (Nelson et al. 1983). Reports in the literature

on the occurrence, geographical distribution and patho-

genicity of these Fusarium species are for the most part

unreliable because of the use of the name F. tricinctum

Corda emend. Snyd. & Hans. (Snyder & Hansen 1945)

which encompasses all four of these species. However,

there is considerable evidence that F. tricinctum sensu

stricto is relatively widespread, but usually at low fre-

quencies, in soils and in association with cereals, includ-

ing wheat, oats, barley and maize, grasses and a number

of other hosts in Europe (Wollenweber & Reinking

1935; Jamalainen 1955; Seemuller 1968; Booth 1971;

Marasas et al. 1979; Domsch et al. 1980; Nirenberg

1981; Ylimaki 1981; Gerlach & Nirenberg 1982; Chel-

kowski et al. 1984) and North America (El-Gholl et al.

1978; Kane et al. 1987). Although F. tricinctum appears

to be relatively common in cold temperate areas of

* Part of a Ph.D. Agric. thesis to be submitted by the first author

to the University of Stellenbosch, Stellenbosch.

** Plant Protection Research Institute, Private Bag X5017, Stel-

lenbosch 7600.

t South African Medical Research Council, P.O. Box 70, Tyger-

berg 7505.

* Department Plant Pathology, University of the Orange Free

State, P.O. Box 339, Bloemfontein 9300.

□ Department of Plant Pathology, University of Stellenbosch,

Stellenbosch 7600.

MS. received: 1987.07.02.

northern Europe, particularly in Finland (Jamalainen

1955; Uoti & Ylimaki 1974; Uoti 1976; Ylimaki

1981), this fungus has also been isolated in Florida in

the USA (El-Gholl et al. 1978). The isolates from leaf-

spots on Hedera helix L. in Florida are the only ones

that have been found to produce the teleomorph ( Gibbe -

rella tricincta El-Gholl, McRitchie, Schoulties & Ridings)

in compatible crosses (El-Gholl et al. 1978, 1979; Cullen

et al. 1983). A dried culture (DAOM 194693) obtained

from CBS 449.67 which was isolated from Calamagros-

tis epigeios (L.). Roth in Berlin, Germany by Seemuller

(1968), has been designated as the neotype for F. tricinc-

tum (Neish 1987).

Two species of the section Sporotrichiella, i.e. F.

poae and F. chlamydosporum are relatively common in

South Africa (Doidge 1938; Doidge 1950; Marasas et

al. 1987). During a recent survey of Fusarium species

in South African soils, three isolates of F. tricinctum

were obtained from Golden Gate National Park (Mara-

sas et al. 1988). These isolates represented the first

record of the occurrence of F. tricinctum in South Africa.

There are also no other authenticated records of this

fungus from the continent of Africa. Subsequently four

additional isolates were obtained. The occurrence, cul-

tural characteristics, morphology and pathogenicity of

these South African isolates of F. tricinctum are describ-

ed and compared with two foreign reference cultures in

this paper.

MATERIALS AND METHODS

Fusarium cultures

The sources of seven South African and two foreign

isolates of F. tricinctum are given in Table 1. All cultures

190

Bothalia 18,2 (1988)

TABLE 1. — Sources of Fusarium tricinctum cultures

* Accession numbers of lyophilised cultures in the collection of the South African Medical Research Council (MRC), Tygerberg,

South Africa.

were initiated from single conidia, lyophilised (Nelson et

al. 1983) and deposited in the culture collection of the

South African Medical Research Council (MRC), P.O.

Box 70, Tygerberg, South Africa. The two foreign refer-

ence cultures were supplied by Dr W. Gerlach, Biologische

Bundesanstalt fur Land- und Forstwirtschaft, Berlin,

Federal Republic of Germany (MRC 1799), and Dr N.E.

El-Gholl, Department of Agriculture, Gainesville, Florida,

USA (MRC 1574). Three isolates from soil debris in

Golden Gate National Park were obtained during a survey

of Fusarium species in South African soils (Marasas et al.

1988). Three additional isolates from soil debris at Lang-

gewens, Malmesbury were obtained by methods identical

to those described by Marasas et al. (1988). A culture

from roots of an annual Medicago sp. (medic) was isolat-

ed as described by Lamprecht etal. (1984).

Cultural and morphological characteristics of all FI tri-

cinctum isolates were determined by incubating cultures

on potato-dextrose agar (PDA, Difco) at 25° C in the

dark and on carnation-leaf agar (CLA, Fisher et al. 1982)

at 20° C under a mixture of white fluorescent and near-

ultraviolet light with a 12 h photoperiod (Nelson et al.

1983). Colony diameters of single-conidial cultures on

PDA incubated in the dark at 25° and 30° C were measured

after 72 h (Burgess & Liddell 1983). Morphological struc-

tures were examined, measured and photographed under

a light microscope.

Culture material for scanning electron microscopy

(SEM) was fixed in 1,5% glutaraldehyde in 0,1 M phos-

phate buffer for 24 h followed by 1 % osmium tetroxide

for 2 h, dehydrated in a graded acetone series, critical

point dried, coated with gold/palladium, and viewed

with an IS1 scanning electron microscope.

Pathogenicity tests

The nine F. tricinctum isolates were tested for patho-

genicity to medics ( Medicago truncatula Gaertn. cv. Jema-

long) and wheat ( Triticum aestivum L. cv. Palmiet) by

means of seed inoculation and soil infestation techniques

described by Lamprecht et al. (1984) and Lamprecht

(1986). In the case of seed inoculation, discs 4 mm diam.,

from 10-d-old water agar (WA, Difco) cultures of each

F. tricinctum isolate, were buried 10 mm deep in steam-

treated soil. Medic seeds, but not the wheat seeds, were

treated with water at 50° C for 30 min. prior to planting.

Six seeds were planted in contact with each disc and

covered with steam- treated soil. There were 60 seeds

per plastic pot (125 mm) and three pots per treatment.

Control seeds were planted in contact with uninoculated

discs. Pots were randomized in a growth chamber at

20° to 22° C. Seedling emergence and damping-off were

followed and recorded after 14 d. Data were analysed by

a one-way analysis of variance and means were compared

using Tukey’s least significant difference (LSD) at the

5% level.

In the case of soil infestation, inoculum of each isolate

of F. tricinctum on a mixture of sand (200 g), bran (10

g) and water (30 ml) was mixed with steam-treated soil

(2,5% , w/w) in 175 mm plastic pots. Control pots con-

tained uninoculated, autoclaved sand-bran mixture. Six-

ty seeds were planted in each pot and there were three

pots per treatment. Pots were randomized and held in a

glasshouse at 15° to 30° C. After six weeks the plants

were removed and disease severity of 10 randomly select-

ed plants per pot scored on a scale from 0 to 5 based on

Bretag (1985) for medics and Kane et al. (1987) for

wheat as follows:

Medics: 0, no visible discolouration of lateral and tap

roots; 1 , slight discolouration of lateral roots, no dis-

colouration of tap root; 2, lateral roots discoloured

brown with lesions, slight discolouration of tap root;

3, necrotic lesions (1 to 5 mm long) on tap root; 4,

necrotic lesions (>5 mm long) on tap root; 5, tap

root completely rotted, plant dead.

Wheat: 0, no visible discolouration of lateral roots and

crown tissue; 1, discolouration of roots and scutellar

node; 2, necrotic lesions on roots and scutellar node;

3, necrotic lesions on roots, scutellar node and sub-

crown intemode, with discolouration extending to

the crown; 4, necrosis of scutellar node and subcrown

intemode with more extensive discolouration of the

crown; 5, extensive necrosis of the crown and stem

base, plant dead.

Bothalia 18,2 (1988)

191

9

e

A

FIGURE 1. — Fusarium tricinctum,

X 710. A, spindle-shaped

and napiform to citriform

micioconidia (MRC 4521);

B, napiform to citriform

microconidia (MRC 4520);

C, spindle-shaped and napi-

form microconidia (MRC

4520); D, monophialide

(MRC 4521); E-F, termi-

nal and intercalary chlamy-

dospores (MRC 4520).

The statistical significance of disease severity indices

was determined by the Kruskal-Wallis one-way analysis

of variance followed by Tukey’s LSD at the 5% level

based on rank averages.

Isolations were made from diseased tissues of re-

presentative plants of both hosts inoculated by both

methods.

RESULTS

Seven isolates of F. tricinctum from three localities in

South Africa (Table 1), were examined. These isolates

were found to be identical to each other and to two

foreign reference cultures of F. tricinctum. The cultural

and morphological characteristics of the South African

isolates of F. tricinctum are described and illustrated

below.

Fusarium tricinctum (Cor da) Saccardo, Sylloge

Fungorum 4: 700 (1886).

See SeemUller (1968), Gerlach & Nirenberg (1982), Nelson

et al. (1983) and Neish (1987) for synonymy.

Colony diameters of single -conidial cultures on PDA

after dark incubation for 3 d are 25—30 mm at 25° C and

12—17 mm at 30° C. Colonies are densely floccose with

irregular margins, pink to carmine red with the reverse

carmine red. Microconidia (Figure 1 A,B,C) are produced

in false heads on monophialides which are formed laterally

on the aerial mycelium or on branched conidiophores,

hyaline, cylindrical and tapered towards the apex (Fig-

ures ID; 3B). Microconidia on PDA and CLA are hyaline,

0— 1-septate, and of two distinct types: fusiform micro-

conidia which are usually produced first (within 14 d)

and abundantly, 10-17 x 3,0— 3,5 /um (Figure 1A, C),

and napiform to citriform microconidia which are pro-

duced later (after 21 d or longer) and sparingly in some

strains, 9—15 x 4—7 jum (Figures 1A, B, C; 3C). Macro-

conidia are produced in orange sporodochia on CLA

after 21 d or longer and are falcate to lunate, with a

pedicellate basal cell and tapered apical cell, 3-5-septate,

25-60 x 3, 0-4, 5 jam (Figures 2A, B, C, D; 3A). Chlamy-

dospores are produced on PDA or CLA after 21 d or

longer, terminal or intercalary, hyaline to pale brown,

smooth to rough-walled, produced singly or in chains

or clumps in the aerial and submerged hyphae, 8—15 jam

diam. (Figures IE, F; 3D).

All nine isolates of F. tricinctum exhibited some de-

gree of pathogenicity to M. truncatula cv. Jemalong as

well as wheat cv. Palmiet (Tables 2 & 3). This fungus

was re-isolated from diseased tissue of inoculated plants

of both hosts, but not from controls. Most isolates were

more pathogenic to M. truncatula than to wheat with

respect to pre- and post-emergence damping-off (Table

2) as well as the severity of root discolouration and ne-

crosis (Table 3). Disease severity indices in medics and

wheat were, however, significantly correlated (r = 0,99,

p <0,05). Isolates from different sources in South Africa

were all weakly to moderately aggressive pathogens and

did not differ much in aggressiveness to either host with

respect to damping-off of seedlings (Table 2). However,

some isolates were significantly (p = 0,05) more aggres-

sive than others with respect to disease severity indices

in medics as well as wheat (Table 3), e.g. isolate MRC

3518 from natural grassland soil at Golden Gate was

significantly more aggressive to both hosts than all other

isolates.

192 Bothalia 18,2 (1988)

TABLE 2. — Pre- and post-emergence damping-off of seedlings of Medicago truncatula cv. Jemalong and Triticum aestivum cv. Palmiet

after seed inoculation with different isolates of Fusarium tricinctum

F. tricinctum Damping-off of seedlings (% )*

isolate

No. (MRC) Medicago truncatula Triticum aestivum

* Each figure based on 1 80 seeds planted per treatment. Means in each column followed by the same letter do not differ significant-

ly (p =0,05).

FIGURE 2. — Macroconidia of Fli-

sarium tricinctum, X 715.

A & B, MRC 3908; C& D,

MRC 4520.

Bothalia 18,2 (1988)

193

FIGURE 3. — Scanning electron-

micrographs of Fusarium

tricinctum. A, macroconi-

dia (MRC 4521), X 1430;

B, monophialides and napi-

form to citriform microco-

nidia (MRC 4521), X 2860;

C, napiform to citriform

microconidia (MRC 4520),

X 1430; D, chlamydospo-

res (MRC 4521), X 1790.

DISCUSSION

The cultural and morphological characteristics of

South African isolates of F. tricinctum were found to be

identical to those of two reference cultures and to agree

with previous descriptions of F. tricinctum (Wollenweber

& Reinking 1935; Booth 1971; Gerlach & Nirenberg

1982; Nelson et al. 1983). The most diagnostic character-

istic of F. tricinctum is the production of napiform to

citriform microconidia on monophialides. It should be

TABLE 3. — Disease development in Medicago truncatula cv.

Jemalong and Triticum aestivum cv. Palmiet plants ex-

posed to soil inoculum of different isolates of Fusarium

tricinctum

F. tricinctum Disease severity index*

isolate

No. (MRC) Medicago truncatula Triticum aestivum

* Each figure represents the mean of 30 plants per treatment.

See Materials and methods for disease severity rating scale.

Means within each column followed by the same letter do not

differ significantly (p =0,05).

noted, however, that these characteristic microconidia

are usually not present in cultures before they are 21 d

old or older, and that they are produced sparingly by

some isolates. In the absence of these microconidia, cul-

tures of F. tricinctum can easily be misidentified as either

F. acuminatum Ell & Ev. or F. reticulatum Mont.

South African isolates of F. tricinctum proved to be

pathogenic to medic as well as wheat seedlings. This is

the first report of the pathogenicity of F. tricinctum to

an annual Medicago sp. (M. truncatula). Isolates of F. tri-

cinctum were weak to moderately aggressive pathogens

of M. truncatula, compared with F avenaceum (Fr.)

Sacc. (Lamprecht et al. 1984; Bretag 1985), and of

wheat, compared with F. graminearum Schwabe Group

1, F. culmorum (W.G. Smith) Sacc. and F. crookwellense

Burgess, Nelson & Toussoun (Van Wyk et al. 1987).

The present results with respect to the pathogenicity to

wheat are in agreement with previous findings (Seemiil-

ler 1968; Uoti 1976; Kane et al. 1987), except that

Seemtiller (1968) found F. tricinctum to be strongly

pathogenic to wheat and to cause damping-off of seed-

lings as well as foot and root rot.

Fusarium tricinctum was not recorded in surveys of

Fusarium species from diseased medics (Lamprecht et al.

1984) and wheat (Van Wyk et al. 1987) in South Africa.

In a recent survey of Fusarium species in South African

soils (Marasas et al. 1988), F. tricinctum was isolated

from one out of 29 soil samples, and only three isolates

of this species out of a total of 2 425 Fusarium isolates

were obtained. On the basis of the available evidence, it

appears that F. tricinctum is a rare fungus in South Africa

194

and is probably not involved as a major pathogen in the

root and crown rot complex of either medics or wheat

in this country. However, in view of the taxonomic con-

fusion surrounding F. tricinctum, the possibility of mis-

identification, and the demonstrated pathogenic poten-

tial of South African isolates of this fungus, researchers

should be aware of the occurrence, distinguishing charac-

teristics, and potential importance of this fungus in South

Africa.

ACKNOWLEDGEMENTS

We thank F.J. Calitz of the Biometry Section, Fruit

and Fruit Technology Research Institute, Stellenbosch

for statistical analyses and A.C. Jaap and M.E. Holtz-

hausen of the Plant Protection Research Institute,

Stellenbosch for competent technical assistance.

REFERENCES

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logical Institute, Kew, England.

BRETAG, T.W. 1985. Fungi associated with root rots of annual

Medicago spp. in Australia. Transactions of the British

Mycological Society 85: 329-334.

BURGESS, L.W. & LIDDELL, C.M. 1983. Laboratory manual

for Fusarium research. Department of Plant Pathology and

Agricultural Entomology, University of Sydney, Sydney.

CHELKOWSKI, J„ VISCONTI, A., SOLFRIZZO, M. & BOT-

TALICO, A. 1984. Formation of mycotoxins by Fusarium

species from cereals in Poland. Phytopathologia Mediter-

ranea 23: 43-46.

CULLEN, D., SMALLEY, E.B. & DIMOND, R.L. 1983. Hetero-

karyosis in Fusarium tricinctum and F. sporotrichioides.

Journal of General Microbiology 129: 3035-3041.

DOIDGE, E.M. 1938. Some South African Fusaria. Bothalia

3: 331-483.

DOIDGE, E.M. 1950. The South African fungi and lichens to

the end of 1945. Bothalia 5: 1-1094.

DOMSCH, K.H., GAMS, W. & ANDERSON, T.H. 1980. Com-

pendium of soil fungi. Vol. 1. Academic Press, London.

EL-GHOLL, N.E., MCRITCHIE, J.J, SCHOULTIES, C.L. &

RIDINGS, W.H. 1978. The identification, induction of

perithecia, and pathogenicity of Gibberella ( Fusarium )

tricincta n. sp. Canadian Journal of Botany 56: 2203-

2206.

EL-GHOLL, N.E., SCHOULTIES, C.L. & RIDINGS, W.H. 1979.

Factors affecting perithecial production in Gibberella (Fu-

sarium) tricincta. Canadian Journal of Botany 57: 2497-

2500.

FISHER, N.L., BURGESS, L.W., TOUSSOUN, T.A. & NELSON,

P.E., 1982. Carnation leaves as a substrate and for preserv-

ing cultures of Fusarium species. Phytopathology 72:

151-153.

GERLACH, W. & NIRENBERG, H. 1982. The genus Fusarium

— a pictorial atlas. Mitteilungen aus der Biologischen Bun-

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desanstalt fur Land- und Forstwirtschaft, Berlin-Dahlem

209: 1-406.

JAMALAINEN, E.A. 1955. Fusarium species causing plant

diseases in Finland. Acta Agricultura Fennica 83: 159-

172.

KANE, R.T., SMILEY, R.W. & SORRELLS, M.E. 1987. Rela-

tive pathogenicity of selected Fusarium species and Micro-

dochium bolleyi to winter wheat in New York. Plant Disease

71: 177-181.

LAMPRECHT, S.C. 1986. A new disease of Medicago truncatula

caused by Cylindrocladium scoparium. Phytophylactica 18:

111-114.

LAMPRECHT, S.C., KNOX-DAVIES, P.S. & MARASAS, W.F.O.

1984. Fusarium spp. associated with diseased root and

crown tissue of annual Medicago spp. Phytophylactica 16:

195-200.

MARASAS, W.F.O. , LEISTNER, L., HOFMANN, G. & ECK ARDT,

C. 1979. Occurrence of toxigenic strains of Fusarium in

maize and barley in Germany. European Journal of Applied

Microbiology and Biotechnology 7: 289-305.

MARASAS, W.F.O., LAMPRECHT, S.C., VAN WYK, P.S. &

ANELICH, R.Y. 1987. Bibliography of Fusarium (Fungi :

Hyphomycetes) in South Africa, 1945-1985. Bothalia 17:

97-104.

MARASAS, W.F.O., BURGESS, L.W., ANELICH, R.Y., LAM-

PRECHT, S.C. & VAN SCHALKWYK, D.J. 1988. Survey

of Fusarium species associated with plant debris in South

African soil. South African Journal of Botany 54: 63-71.

NEISH, G.A. 1987. Neotypification of Fusarium tricinctum.

Canadian Journal of Botany 65: 589-591.

NELSON, P.E., TOUSSOUN, T.A. & MARASAS, W.F.O. 1983.

Fusarium species. An illustrated manual for identification.

Pennsylvania State University Press, University Park.

NIRENBERG, H. 1981. A simplified method for identifying

Fusarium spp. occurring on wheat. Canadian Journal of

Botany 59:1599-1609.

SEEMULLER, E. 1968. Untersuchungen iiber die morphologische

und biologische Differenzierung in der Fusarium Sektion

Sporotrichiella. Mitteilungen aus der Biologischen Bundes-

anstalt fur Land- und Forstwirtschaft, Berlin-Dahlem 127:

1-93.

SNYDER, W.C. & HANSEN, H.N. 1945. The species concept in

Fusarium with reference to Discolor and other sections.

American Journal of Botany 32: 657-666.

UOTI, J. 1976. The effect of five Fusarium species on the growth

and development of spring wheat and barley. Annales Agri-

culture Fenniae 15: 254-262.

UOTI, J. & YLIMAKI, A. 1974. The occurrence of Fusarium

species in cereal grain in Finland. Annales Agriculturae Fen-

niae 13: 5-17.

VAN WYK, P.S., LOS, O., PAUER, G.D.C. & MARASAS, W.F.O.

1987. Geographic distribution and pathogenicity of Fusa-

rium species associated with crown rot of wheat in the

Orange Free State, South Africa. Phytophylactica 19: 27 1 —

274.

WOLLENWEBER, H.W. & REINKING, O.A. 1935. DieFusarien.

Ihre Beschreibung, Schadwirkung und Bekampfung. Paul

Parey, Berlin.

YLIMAKI, A. 1981. The mycoflora of cereal seeds and some

feedstuffs. Annales Agriculturae Fenniae 20: 74-88.

Bothalia 18,2: 195-209 (1988)

Leaf anatomy of the South African Danthonieae (Poaceae). XVII. The

genus Chaetobromus

R.P. ELLIS*

Keywords: Chaetobromus, Danthonieae, leaf anatomy, Poaceae

ABSTRACT

The leaf anatomy of Chaetobromus Nees is described and illustrated. The genus has non-Kranz anatomy but

with a low lateral cell count. The abaxial epidermis is not pooid as microhairs are present, the silica bodies are

dumbbell-shaped and the stomatal subsidiaries are dome-shaped. From the sample of 33 randomly selected speci-

mens from throughout the distribution range of the genus it was possible to identify four anatomical groups,

each associated with a specific habitat and differing in vegetative morphology. These intergrade in a reticulate

pattern and do not constitute discrete entities. It appears that the genus is monospecific with four poorly separated

subspecific taxa.

UITTREKSEL

Die blaaranatomie van Chaetobromus Nees word beskryf en geillustreer. Die genus het nie-Kranzanatomie

maar met ’n lae laterale seltelling. Die abaksiale epidermis is nie pooied nie aangesien mikrohare aanwesig is, die

silikaliggame murgbeenvormig en die hulpselle van die huidmondjies koepelvormig is. Uit ’n monster van 33 ek-

semplare wat willekeurig vanuit die hele verspreidingsgebied van die genus gekies is, kon vier anatomiese groepe

geidentifiseer word, elk met ’n spesifleke habitat geassosieer, en verskillend wat vegetatiewe morfologie betref.

Hierdie groepe vloei ineen in ’n netvormige patroon en vorm nie afsonderlike entiteite nie. Dit blyk dat die genus

monospesifiek is met vier swak omgrensende subspesifieke taksons.

INTRODUCTION

Chaetobromus Nees is a small genus in which four

closely allied species have been described. However, they

are very difficult to distinguish from one another and

must be regarded as provisional until the genus has been

studied in greater detail. Nowadays only three specific

names are generally accepted (Clayton & Renvoize

1986). These are C. dregeanus Nees, C. involucratus

(Schrad.) Nees and C. schraderi Stapf. The name C.

schlechteri Stapf, that of the fourth species described,

has fallen into disuse although it is correct (Smook &

Gibbs Russell 1985). Chippindall (1955) does not con-

sider C. schlechteri to be distinct from C. dregeanus.

Chaetobromus is indigenous to southern Africa with

the centre of distribution in the western Cape, Namaqua-

land and southern South West Africa/Namibia. The genus

occurs mainly in the Strandveld, the Namaqualand Coast

Belt of the Succulent Karoo and the Namaqualand Broken

Veld (Acocks 1975). All these veld types are at low al-

titudes, with sandy soil and with winter rainfall decreas-

ing northwards. It is also occasionally found further

south in the southern Strandveld as far south as Table

Bay. It is of interest that Chaetobromus also occurs at

higher altitudes in the Western Mountain Karoo from

Loeriesfontein southwards to Sutherland. It is rare in

this latter veld type but this appears to be due to over-

grazing.

The leaf anatomy of Chaetobromus has received little

attention in the literature. De Wet (1956) reported that

* Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

MS. received: 1987.10.20.

the epidermis is panicoid, with linear microhairs and

dumbbell-shaped silica bodies, and the mesophyll is

‘festucoid’ with a poorly differentiated outer bundle

sheath and uniformly distributed chlorenchyma. Watson

et al. (1986) give further anatomical details. They con-

sider the silica bodies to be intermediate between the

panicoid and the pooid type. Stomata are present with

guard cells flush or overlapping the interstomatals and

the subsidiary cells are either parallel-sided or low dome-

shaped — either panicoid or pooid. These authors also

did not observe abaxial microhairs. They consider the

genus to be C3 although the lateral cell count is rather

low. The midrib is structurally indistinguishable from

the other larger bundles. All bundles have wide adaxial

bundle sheath extensions. The bulliform cells occur in

very large, simple, fan-shaped groups without associated

colourless cells. The present paper will describe and il-

lustrate the anatomy of the genus based on a representa-

tive sample and will compare this anatomy with the

findings of the above authors.

MATERIALS AND METHODS

Leaf blade segments of Chaetobromus plants were

freshly fixed in the field in FAA (Johansen 1940). Her-

barium voucher specimens from the same plants were

taken for determination by the National Herbarium

(PRE) where they are housed.

Leaf transverse sections, 10 nm thick, were prepared

after desilicification in hydrofluoric acid (Breakwell

1914), dehydration using the method of Feder& O’Brien

(1968) and embedding in Tissue Prep (Fischer Scientific).

These sections were stained in safranin and fast green

(Johansen 1940). The manual scraping method of Met-

calfe (1960) was used to prepare scrapes of the abaxial

196

Bothalia 18,2 (1988)

FIGURE 1. — Leaf blade anatomy of Chaetobromus specimens representative of Group 1 as seen in transverse section. A, outline

showing sectioned macrohairs, Ellis 2182, X 100. B, slightly inrolled outline, Ellis 2187, X 100. C-D, Ellis 2184: C, leaf

• outline, X 100; D, anatomical detail of chlorenchyma and vascular bundles with abaxial macrohairs, X 250. E-F, Ellis

21 76: E, outline of margin, X 100; F, detail of chlorenchyma with low lateral cell count, X 250.

leaf epidermis. The anatomical structure was recorded

photographically using a Reicherdt Univar microscope

and Ilford Pan F Film (50 ASA).

The standardized terminology of Ellis (1976, 1979)

was used for the anatomical descriptions. The following

abbreviations will be used in these descriptions:

vb/s — vascular bundle/s

l’vb/s — first order vascular bundle/s

2’vb/s — second order vascular bundle/s

3’vb/s — third order vascular bundle/s

ibs — inner bundle sheath; mestome sheath

obs — outer bundle sheath; parenchyma sheath

Identification of the voucher specimens to species

level proved to be very difficult and unsatisfactory. This

is understandable as the genus is in need of revision

(Chippindall 1955), the species concepts are vague, and

species are indistinct and intergrade, with many inter-

mediate specimens being present. It has, therefore, been

decided to consider this genus at the generic level only.

Separate descriptions of the species will not be given due

to uncertainty of the exact identity of a significant

proportion of the voucher specimens. The anatomical

description of the genus given is inclusive of the entire

sample of 33 specimens used in this study.

It was, nevertheless, possible to sort the material

studied into four informal groups using anatomical, eco-

logical and vegetative morphological criteria. These

informal groups appear to correspond reasonably well

with the species described in the genus and they will be

compared and contrasted in an attempt to ascertain

whether they represent distinct species. In the citation

of the specimens examined these informal groups are

used together with their probable specific equivalent.

Asterisks indicate cases where there is a discrepancy

between the herbarium determination (in brackets) and

the grouping used here.

Bothalia 18,2 (1988)

197

FIGURE 2. — Abaxial epidermal structure of Chaetobromus specimens of Group 1 as seen in surface view. A-B, Ellis 21 75; A,

macrohairs, many filled with air, X 160; B, detail of macrohair bases and silica bodies, X 250. C, pubescence of leaf sur-

face, phase contrast, Ellis 2187, X 100. D, macrohairs, silica bodies and intercostal microhairs, Ellis 2184, X 250.

Specimens examined

Group 1 . Chaetobromus involucratus

CAPE. — 2816 (Oranjemund): Richtersveld, Beesbank (-BC),

Ellis 50 71 ; Beauvallon (-DA), Ellis 21 75*, 21 76 * (C. schraderi).

2916 (Port Nolloth): Port Nolloth (-BD), Ellis 2182, 2183,

2184, 2185, 2187. 3017 (Hondeklipbaai): Hondeklip Bay (-AD),

Ellis 5344, 5345. 3217 (Vredenburg): St Helena Bay, Stompneus-

baai (-DB), Ellis 703* (C. schraderi).

Group 2. Chaetobromus schraderi

CAPE. — 2816 (Oranjemund): Richtersveld, Beesbank (-BC),

Ellis 5073. 2917 (Springbok); Steinkopf (-BD), Ellis 2164, 2165,

2166; Spektakelberg, Sandhoogte Pass (-DA), Ellis 2191, 2192,

2193, 2194. 3018 (Kamiesberg): 43 km N of Bitterfontein

(-CA), Ellis 1730* (C. dregeanus ); Bitterfontein (-CC ), Ellis

2206. 3118 (Van Rhynsdorp): 64 km N of Van Rhynsdorp

(-BC), Ellis 1 733* (C. dregeanus).

Group 3. Chaetobromus dregeanus (lowland form)

CAPE. — 3218 (Clanwilliam): Elands Bay (-AD), Ellis 1691.

3317 (Saldanha): Saldanha Bay (-BB), Ellis 1696. 3318 (Cape

Town): Yzerfontein (-AC), Ellis 692; Blouberg Strand (-CD),

Ellis 2357, 2358.

Group 4. Chaetobromus dregeanus (mountain form)

CAPE. — 3019 (Loeriesfontein): Loeriesfontein at junction

of Nieuwoudtville and Granaatboskolk roads (-CD), Ellis 2422;

103 km from Brandvlei on road to Loeriesfontein (-DC), Ellis

2423, 2424. 3118 (Van Rhynsdorp): 10 km S of Bitterfontein

(-AB), Ellis 2426. 3119 (Calvinia): Van Rhyns Pass (-AC),

Ellis 2457; Botterkloof Pass between Calvinia and Clanwilliam

(-CD), Ellis 2450. 3220 (Sutherland): Sutherland (-BC), Ellis

2467.

LEAF ANATOMY OF THE GENUS CHAETOBROMUS

Leaf in transverse section

Outline : open, expanded, flat (Figures 3A— C, 5A &

B) to slightly inrolled (Figures 1A— C, 5D) or narrow,

infolded (Figure 7A, C, G & H) or inrolled (Figure 7E);

leaf thickness 230—550 pm. Ribs and furrows : slight

(Figure 3 A— D) to medium (Figure 1A— F) rounded

adaxial ribs overlie l’vbs and 2’vbs; furrows shallow and

wide (Figure 3A-D) to narrow, cleft-like (Figure 7E

& F); furrows may be present over 3’vbs; slight abaxial

ribs and furrows sometimes present (Figures 1 A-F, 7H),

usually not developed. Median vascular bundle : not

structurally distinct from lateral l’vbs. Vascular bundle

arrangement ; tends to be variable and rather irregular

although a basic pattern is discernible; 5, 7, 9 or 1 1 l’vbs

in leaf section; one 2’vb between consecutive l’vbs al-

though this sometimes varies; one 3’vb between adjacent

l’vbs and 2’vbs but may be absent but then usually only

on one side of each 2’vb (Figures IF, 3E & 5C). All vbs

situated in centre of blade. Vascular bundle description;

l’vbs round to elliptical in shape; phloem adjoins ibs;

metaxylem vessels very narrow, diameters much less

than those of obs cells (Figures ID & F, 3D & F, 5E &

F); 2’vbs and 3’vbs round to elliptical; xylem and

phloem distinguishable. Vascular bundle sheaths; double;

obs round to elliptical; prominent adaxial and abaxial

multiseriate parenchymatous extensions extend to the

198

Bothalia 18,2 (1988)

upper and lower epidermis from all l’vbs (Figures ID &

F, 3D & F, 5E & F, 7B, D & F); 2’vbs with narrower

adaxial extensions, being bi- or uniseriate; 3’vbs with

irregular extensions or extensions absent; obs cells in-

flated, round, the cells smaller than the mesophyll cells;

walls thin; either translucent, without chloroplasts

(Figures 3D & 7F) or with small chloroplasts, fewer than

in the chlorenchyma cells (Figures ID & F, 3E & F, 5C

& E); ibs entire, of small lignified cells, those opposite

the phloem with the inner tangential and radial walls

thickened (Figures 3D & 5C). Sclerenchyma: minute

adaxial and slightly larger abaxial strands associated with

the l’vbs and 2’vbs; no sclerenchyma adjacent to the

3’vbs; small sclerenchyma caps in the margins; cell walls

not lignified. Chlorenchyma: irregular (Figure 5E & F)

to tending to radiate condition (Figures ID & F, 2D— F);

more than one layer of radiating cells; lateral cell count

low, from 3—5 cells; continuous between bundles;

chlorenchyma cells large, angular and equidimensional

(Figure ID & F) to elongated (Figure 3D— F); often

nucleate and may be refractive under interference con-

trast (Figure 3F); no arm cells or fusoids; intercellular

air spaces large and frequent. Colourless cells: absent ex-

cept in the bundle sheath extensions. Adaxial epidermal

cells: small to medium-sized bulliform groups at bases

of furrows (Figures ID & E, 3D&F, 5E&F, 7F); occupy

less than V4 leaf thickness; central cell often largest and

may be shield-shaped; cuticle very thin; epidermal ap-

pendages usually absent but macrohairs (Figure lA)and

prickles (Figure 5E)may be present; no papillae. Abaxial

epidermal cells: bulliform cells absent; cuticle very thin

or slightly thickened; macrohairs present (Figure 1A, C

& D) or absent; no prickles or papillae.

Abaxial epidermis in surface view

Intercostal long cells: elongate rectangular (Figures

2B & D, 4C & E, 8F) or fusiform (Figures 4D, 6A— D,

8A— D); walls straight (Figures 2A-D, 4A— E) to slight-

ly sinuous (Figures 6A— F, 8A— F); cell shape and size

consistent across individual intercostal zones; long cells

adjoin one another or are separated by short cells; no

bulliform cells. Stomata: low dome-shaped (Figures 2B

& D, 4C & E) to dome-shaped (Figures 6B, D, F, 8A-F)

subsidiary cells; occurring throughout intercostal zones

FIGURE 3. — Transectional leaf anatomy of Group 2 representatives of Chaetobromus. A, expanded, flat outline, Ellis 2165,

X 100. B, leaf margin, Ellis 2164, X 100. C-D, Ellis 2166: C, outline, X 100; D, chlorenchyma cell arrangement and vas-

cular bundles with parenchymatous extensions, X 250. E— F, Ellis 2191: E, vascular bundle arrangement, X 160; F, detail

of chloroplasts in chlorenchyma and bundle sheath cells, interference contrast, X 250.

Bothalia 18,2 (1988)

199

or restricted to two or four rows; stomata in files separat-

ed by single interstomatal long cell, this either short or

elongated, more than three times longer than wide;

tend to be overarched by inflated long cells (Figure 9F).

Intercostal short cells: presence variable, either absent

or irregularly present to present between all long cells;

either single or paired; tall and narrow in shape. Papillae:

absent. Prickles: absent except for very small hooks

which are rarely present (Figure 6D ). Microhairs : present

or absent; elongated, basal and distal cells about equal

in length (Figures 4C, 9H) or distal cell much shorter

(Figure 9D); hairs short, about the same length as the

stomata; distal cell deciduous and thin-walled. Macro-

hairs: usually absent but may be present (Figures 2A— D,

9A, B); unicellular, hard, stiff and pointed; base con-

stricted and superficial, not associated with specialized

raised epidermal cells (Figures 9B, 10); appressed to leaf

surface and visible in transections (Figure 1A, C & D);

length up to 120 /am; tend to be very numerous; lumens

often filled with air (Figure 2A & B); intercostal. Silica

bodies : horizontally elongated, variable dumbbell-shaped

(Figures 2B, 4E, 6B & D, 8B, D & F); rarely nodular

(Figure 6F); exclusively costal; granules may be present.

Adaxial epidermis in surface view

Differs from abaxial epidermis in having shorter inter-

costal long cells, costal prickles with elongated barbs are

common (Figure 9C & G) and numerous microhairs are

present adjacent to the costal zones (Figure 4F); the

microhairs are larger and more common than on the

abaxial surface; the distal cell may also be rounded and

not tapering (Figure 9H). These differences in frequency

of occurrence, size and shape of the adaxial and abaxial

microhairs on the same leaf are unusual.

FIGURE 4. — A-E, abaxial epidermis of Group 2 Chaetobromus specimens: A-C, Ellis 2191: A, zonation pattern, X 160; B,

intercostal zone showing absence of macrohairs, X 250; C, detail of subsidiary cell shape, intercostal long cells and micro-

hairs, X 400. D, fusiform intercostal long cells, Ellis 2164, X 250. E, interference contrast showing microhairs, Ellis 1 730 ,

X 250. F, adaxial epidermis showing prickle hairs and microhairs, Ellis 2206, X 400.

200

Bothalia 18,2 (1988)

FIGURE 5. — Transverse sections of Group 3 Chaetobromus specimens. A, outline, Ellis 2358, X 100. B, outline and leaf margin,

Ellis 1691, X 100. C, anatomical detail of chlorenchyma and vascular bundles, Ellis 2357, X 250. D-F, Ellis 1696: D,

slightly inrolled outline, X 100; E, chloroplast distribution in bundle sheath and chlorenchyma cells, X 250; F, interference

contrast showing chloroplast and bundle sheath cell shape, X 250.

DISCUSSION

Subfamilial and tribal classification

The results of this study confirm and expand the find-

ings of De Wet (1956) and Watson etal. (1986) who re-

ported a ‘festucoid’ transectional anatomy and a panicoid

abaxial epidermis for Chaetobromus . The abaxial epider-

mis is not pooid in several respects such as the possession

of microhairs, dumbbell -shaped silica bodies and dome-

shaped stomata, and as a result, pooid relationships can

be discounted. Microhairs are unknown in pooid grasses

and dumbbell-shaped silica bodies are very rare (Watson

et al. 1985). Watson et al. (1986) did not observe abaxial

microhairs on Chaetobromus but their presence is con-

firmed here although they are rare and appear to be absent

on those specimens assigned to groups 3 and 4. Panicoid

type dumbbell-shaped silica bodies are common although

the nodular type of pooid body was sometimes present.

This confirms the observations of Watson et al. (1986).

The pooid type of stomata, in which the guard cells over-

lap the interstomatal cells (Watson & Johnston 1978),

do not appear to occur in Chaetobromus as reported by

Watson et al. (1986). Instead the guard cells are flush

with the interstomatals and the subsidiary cells are dome-

shaped. This is typical of the panicoid type and not pooid

where the subsidiaries are typically parallel-sided.

The transectional anatomy of Chaetobromus, on the

other hand, can be considered to be of the pooid type.

The outer bundle sheath cells are non-Kranz and do

not possess specialized chloroplasts and have thin cell

walls. The chlorenchyma cells are large parenchymatous

cells usually diffusely arranged, often tending to the

radiate condition. The genus is undoubtedly C3 as sug-

gested by Watson et al. (1986) although this has yet to

be confirmed physiologically.

This combination of pooid transectional anatomy

with a panicoid type of epidermis is typical of the Arun-

dinoideae. The leaf anatomy, therefore, supports arundi-

noid affinities for Chaetobromus and is in full agreement

with the classification of the genus in the Arundineae

(Clayton & Renvoize 1986).

Certain aspects of the chlorenchyma of Chaetobromus

deserve further comment. As noted by Watson et al.

(1986), the lateral cell count is rather low, from 3-5

Bothalia 18,2 (1988)

201

FIGURE 6. — Abaxial epidermal structure of Chaetobromus specimens of Group 3. A-B, Ellis 2357: A, costal and intercostal

zone distribution, X 160; B, fusiform long cells and differential staining of intercostal long cells, X 250. C-D, Ellis 2358:

C, costal and intercostal zones, X 160; D, costal and intercostal silica bodies and short intercostal hooks, X 250. E, epider-

mal pattern, Ellis 1696, X 160. F, subsidiary cell and silica body shape, Ellis 692, X 400.

cells, which is unusual in a C3 plant. The chlorenchyma

arrangement may also tend to the radial type of arrange-

ment although more than one layer of radiating cells is

always present. This chlorenchyma arrangement re-

sembles that of Dregeochloa pumila (Nees) Conert (Ellis

1977), another danthonioid from the winter rainfall

coastal areas of the southern Namib Desert. Merxmuel-

lera rangei (Pilg.) Conert appears to represent the ex-

treme of this type of development (EUis 1982). The

vegetation of this area is dominated by succulents and

the chlorenchyma of these associated grasses is reminis-

cent of that of CAM plants, particularly the large size

of the cells, their particular radial type of arrangement,

the persistent nuclei, large vacuoles and the refractive

nature of the chloroplasts under interference contrast

illumination. The CAM photosynthetic pathway is un-

known in the Poaceae but, if it does occur, these grasses

appear to be the most likely candidates. Their photo-

synthetic physiology deserves investigation as it may

demonstrate the presence of CAM in the grass family.

Differences between the species of Chaetobromus

The basic anatomical differences between the four

species or groups in the genus are listed in Table 1.

These are usually not discrete differences and they

often intergrade between the groups. However, they do

give an indication of the anatomical tendencies evident

within the genus.

TABLE 1. — The main anatomical differences separating the infrageneric taxa of Chaetobromus

202

Bothalia 18,2 (1988)

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Group 1 . Chaetobromus involucratus

Anatomy

The leaf blades are expanded but tend to roll inwards

from both margins when desiccated. The well developed

adaxial ribs and steep-sided furrows (Figure ID & F)

may be associated with this inrolling mechanism. The

blades are of intermediate thickness and width and usually

contain 7 first order vascular bundles. The chlorenchyma

cells are angular to equidimensional and not elongated

as in Group 2.

The abaxial macrohairs are diagnostic for this group

(Figures 2A-D, 9A, B, 10). Rectangular long cells with-

out sinuous walls and the presence of microhairs (al-

though rare) characterize the epidermis of this group.

These microhairs have tapering distal cells which may be

equal to or much shorter than the basal cell (Figure 9D).

The diagnostic abaxial macrohairs are not always pro-

duced on all leaves or on all plants. Chippindall (1955)

considers these hairs to be deciduous as the older leaves

appear to become glabrous. The epidermal anatomy con-

firms that their bases are only superficial and are not

inserted between the epidermal cells. Instead they are

attached to a small, raised intercostal short cell (Figure

10) which possibly acts as a hinge mechanism. On leaves

with few or no hairs, however, there is no evidence of

these short cells, or of the slight disruption in the inter-

costal long cell pattern caused by these bases (Figure 2B).

This evidence appears to indicate that these hairs are not

deciduous but that their presence is variable, either tem-

porally or spatially.

The function of these hairs is unknown but, as macro-

hairs or prickles also occur on other perennial grasses

from this fog belt, they may help in trapping moisture

from the regular sea mists. Examples are Dregeochloa

pumila (Ellis 1977) and Centropodia glauca (Nees) T.A.

Cope (Ellis 1984).

Morphology and ecology’

Found in loose windblown sand and unstabilized

coastal dunes of the Strandveld of Namaqualand. Particu-

larly common in the north from Hondeklip Bay to Alexan-

der Bay but extending as far southward as St Helena Bay

( Ellis 703). The rainfall of this region is very low, 50—

150 mm per annum mainly in winter, but it is subject

to frequent sea mists. The vegetation is a sparse succulent

scrub and the substrate is loose and sandy.

Populations of this taxon are exceedingly variable in

vegetative morphology, with each individual plant dif-

fering in size, foliage colour and growth form. Plants are

generally erect tufts up to 0,3 m high with the leaves

being borne all the way up the culms. The inflorescences

are not well exserted. Sometimes the plants are basally

tufted but this appears to be in response to grazing. Rhi-

zomes are not prominent except in these basally tufted

specimens. The roots often are enclosed in a prominent

rhizosphere of sand grains.

The leaf blades are expanded to slightly inrolled and

are densely hairy with soft, silky, appressed hairs on the

203

abaxial or both surfaces. This is particularly the case

when the leaves are young but, according to Chippindall

(1955), they become almost glabrous when older. This

is, however, not supported by the available anatomical

evidence. There is also a definite decrease in the degree

of pubescence southwards from the Orange River and

the specimen from St Helena Bay has only a few marginal

hairs. However, specimens from the north may also have

few hairs.

The Group 1 specimens grade into the Group 3 type

from St Helena Bay southwards and their distribution

is sympatric between St Helena and Elands Bay. South

of this area sea mists decrease in relative importance and

frequency and this may account for the loss of the

macrohairs.

The morphology of the Group 1 plants conforms

closely to the description of C. involucratus (Chippin-

dall 1955) and most specimens assigned to this group

were identified as such. However, a few were identified

as C. schraderi because these usually had very few macro-

hairs as in Ellis 703 and 2176 (Figure IE & F) but the

blade of Ellis 21 75, which was examined anatomically,

had a dense pubescence (Figure 2 A & B).

Group 1 is therefore virtually synonymous with

C. involucratus and is reasonably distinct from the other

taxa. It is distinguished morphologically by the presence

of the diagnostic hairs and it also occupies a specialized

habitat. However, it does intergrade with both Groups

2 and 3, both in morphology and ecology, and they can-

not always be clearly separated.

Group 2. Oiaetobromus schraderi

Anatomy

All specimens have expanded but flat leaf blades

which appear to fold in response to moisture stress. This

contrasts to the inrolling of the leaves of Groups 1 & 3.

The adaxial ribs and furrows are also poorly developed

(Figures 3E & F) and this may be associated with the

infolding rather than an inrolling mechanism. The leaf

blades are thicker than in any of the other groups and

may be up to 550 pm thick. The leaves also tend to be

wider with an average of 9 first order bundles. The chlo-

renchyma is distinctive with large, elongated or rounded

cells arranged in a radiate manner; the nuclei tend to be

persistent and the chloroplasts may be refractive. This

chlorenchyma structure resembles that of some liliaceous

CAM plants and Group 2 plants may exhibit modifica-

tions of the C3 photosynthetic pathway.

Macrohairs are absent and the long cells have straight

walls and are usually rectangular as in Group 1. However,

they do vary and may be fusiform in shape (Figure 4D).

Microhairs are present although uncommon and the distal

cell does not taper to a pointed apex (Figure 9H).

The transverse sections of the Group 2 plants are dis-

tinctive but the abaxial epidermis resembles that of Group

1 more than that of the other two anatomical groups.

The main differences between Groups 1 and 2 are the

absence of macrohairs and the shape of the microhairs.

204

Bothalia 18,2 (1988)

FIGURE 7. — Transections of Group 4 specimens of Chaetobromus . A, narrow, infolded leaf blade, Ellis 2424, X 100. B, vascu-

lar bundle arrangement, interference contrast, Ellis 2423, X 160. C-D, Ellis 2467 : C, infolded leaf, X 100; D, detail of

chloroplasts in chlorenchyma and bundle sheath cells, X 250. E— F, Ellis 2450: E, inrolled, setaceous outline, X 160; F,

vascular bundles showing low lateral cell count, X 250. G, outline of specimen from mountain fynbos on sandstone, Ellis

2457, X 100. H, outline of specimen from Succulent Karoo, Ellis 2426, X 100.

Bothalia 18,2 (1988) 205

FIGURE 8. — Abaxial epidermis of Group 4 Chaetobromus specimens. A-B, Ellis 2424 : A, epidermal zonation, X 160; B, de-

tail of fusiform long cells and stomata, interference contrast, X 250. C-D, Ellis 2457 : C, interference contrast showing

numerous stomata in regular files, X 160; D, detail of stomata, intercostal long cells and costal cells (note absence of mi-

crohairs), X 250. E, epidermal pattern with differential staining, Ellis 2467, X 160. F, stomata and silica bodies, inter-

ference contrast, Ellis 2422, X 250.

Morphology and ecology

This group is found in the Namaqualand Coast Belt

of the Succulent Karoo and in Namaqualand Broken

Veld. It appears to prefer heavier stony soils, often be-

tween granite rocks in the mountains of Namaqualand.

It also occurs on the sand of the coastal plain but not on

unstabilized sand, the habitat characteristic of Group 1.

The specimens Ellis 5071 (Group 1) and Ellis 5073

(Group 2) are informative in this regard as they were

found growing in close proximity to each other but

differ considerably in epidermal structure (Figure 9A-

H). Ellis 5073 was rooted in gravelly soil between granite

boulders which had subsequently been covered by fine,

loose wind-blown sand in which Ellis 5071 was growing.

Group 2 plants are most common along roadside verges

where they are protected from grazing.

The distribution area extends from the Richtersveld

in the north to the Vanrhynsdorp District in the south.

This area receives a sparse winter rainfall varying between

50-300 mm per annum with fog decreasing in importance.

The vegetation is dominated by succulents.

Populations of Group 2 plants are very variable with

individual plants exhibiting different growth forms and

vegetative morphology. Generally they are robust, tall

plants up to 0,9 m tall. The leaves are not basally con-

centrated but are cauline. These leaves are long and ex-

panded, up to 10 mm wide and glabrous. The leaf sheaths

may be hairy and the adaxial surface has short prickles

(Figure 4F). The rhizomes are short but well developed

and inclined upwards resulting in loosely caespitose

plants. The roots are tough and corky and a rhizosphere

is seldom developed.

206

Bothalia 18,2 (1988)

Bothalia 18,2 (1988)

207

The name applied to most specimens of this group is

C. schraderi although two specimens were identified as

C. dregeanus. Morphologically and ecologically this

group is not well separated from C. involucratus (Group

1) and appears to be an inland segregate of that species.

Chippindall (1955) considered C. schraderi to be a sand

dune species found south of Saldanha Bay to Table Bay.

This distribution corresponds with the Group 3 speci-

mens of the present study, all of which were determined

as C. dregeanus. The possibility therefore exists that

these names have been misapplied.

Although no consistent and reliable vegetative and

ecological diagnostic characters appear to distinguish

C. involucratus (Group 1) from C. schraderi (Group 2),

leaf anatomy reveals differences which may justify the

upholding of these two taxa.

Group 3. Chaetobromus dregeanus (lowland form)

Anatomy

The leaf blades are flattened to slightly inrolled with

adaxial ribs and furrows and are generally thinner than

in either Group 1 or 2. The chlorenchyma cells resemble

those of Group 1, being angular and equidimensional

(Figures 5C & F) and not as large and elongated as in

Group 2. The transectional anatomy is, therefore, very

similar to that of Group 1 .

The abaxial epidermis, on the other hand, differs

considerably from that of Group 1. Macrohairs and

microhairs are absent and the long cells are sinuous.

This epidermal structure resembles that of Group 4.

Morphology and ecology

Only a small sample of this taxon was studied but all

specimens were collected in the southern Strandveld,

from Elands Bay southwards to Table Bay, in the Dense

Strandveld Scrub variation of this veld type (Acocks

1975). Winter rainfall is higher (200—300 mm) than

further north and fog is infrequent. The vegetation con-

sists of a dense, dwarf, semi-succulent scrub, about 1 m

tall. The sand is stabilized, the dunes being well vegetated

and the soil has a high humus content. This is clearly

evident on the roots which are black in colour.

The plants form loose, diffuse tufts up to 0,4 m high.

The culms are not basally concentrated but dispersed

and linked by rhizomes or even stolons up to 0,25 m

long. Scattered culms, therefore, protrude through the

dense scrubby vegetation. The leaves are borne up the

culms and are expanded to slightly inrolled but glabrous.

This group appears to represent a southward extension

of C. involucratus (Group 1) which has become adapted

to the absence of fog, a denser vegetation and stabilized,

FIGURE 10. — Base and epidermal attachment of abaxial macro-

hair of Group 1, Chaetobromus involucratus, Ellis 5071,

X 660.

humic sand. Unexpectedly the identification of the

voucher specimens does not reflect this relationship as

they were all assigned to C. dregeanus, a species which

does not appear to be closely linked to C. involucratus.

Judging by the identification, Group 3 is therefore con-

sidered to be merely a lowland form of C. dregeanus.

This view, however, is in conflict with the evidence pre-

sented here. Chippindall (1955) considered C. schraderi

to occur in these southern Strandveld areas and this

taxon may, therefore, actually be C. schraderi. The typi-

fication of the names applied to these Chaetobromus taxa

is required to resolve these nomenclatural difficulties.

Group 4. Chaetobromus dregeanus (mountain form)

Anatomy

The transectional anatomy of this group is diagnostic,

the leaves being narrow and strongly infolded or inrolled.

In some cases this reduction resembles the permanently

infolded, setaceous condition (Figure 7E). Associated

with this narrowing of the leaf blade is a reduction in

the number of first order vascular bundles to 5 and the

loss of the second order bundles. The cleft-like adaxial

furrows may also result from the extreme infolding ex-

hibited by some specimens of this group (Figure 7E &

F).

The epidermis bears a close resemblance to that of

Group 3 due to the sinuous long cell walls and the ab-

sence of microhairs and macrohairs.

Morphology and ecology

This group includes all those Chaetobromus speci-

mens from the inland, mountainous areas of the western

FIGURE 9. — SEM micrographs showing the abaxial and adaxial epidermal structure of specimens of Chaetobromus involucratus

and C. schraderi collected at the same locality. A-D, Ellis 5071, Group 1: A, abaxial epidermis showing dense pubescence of

intercostal macrohairs, X 66; B, detail of macrohairs showing superficial bases, X 204; C, adaxial epidermis with prickle hairs

on ribs which differ in size and base structure from the abaxial macrohairs, X 204; D, abaxial microhair with very short,

tapering distal cell, X 900. E-H, Ellis 5073, Group 2: E, abaxial epidermis lacking macrohairs, X 66; F, detail of inflated

intercostal long cells overarching the slightly sunken stomata, X 204; G, adaxial epidermal ribs with prickle hairs with short

barbs, X 204; H, adaxial microhair showing basal and distal cells of equal length and the distal cell with a blunt apex, X 1320.

208

Bothalia 18,2 (1988)

Cape. Most representatives of this group were collected

in the Western Mountain Karoo, particularly in the lower

or semi-succulent form of this veld type (Acocks 1975)

from the Loeriesfontein, Calvinia and Sutherland Dis-

tricts. These areas range in altitude from 600-1000 m

above sea level and receive 150 mm and less of rain,

mostly in winter. They are dry, heavily overgrazed areas

with a sparse karroid vegetation. Nowadays Chaeto-

bromus is only found in protected sites between rocks

and under bushes but it may formerly have been much

more common, extending further eastwards as far as

Fraserburg and even Hanover (Chippindall 1955).

A few other specimens with narrow, infolded leaves

have also been included in this group although they differ

slightly in leaf anatomy and occur in other veld types.

The sample studied is too small to draw meaningful con-

clusions but these may deserve recognition as separate

groups. Ellis 2457 is the only specimen studied which

originated in Mountain Fynbos on soil derived from Table

Mountain Sandstone. The anatomy differs little from

that characteristic of this group although abaxial stomata

are particularly common (Figures 7G, 8C & D). Ellis

2426 is the other specimen assigned to this group on

anatomical grounds but it was collected in the Succulent

Karoo on very stony soil. This specimen is characterized

by having well developed sclerenchyma strands and

adaxial ribs and furrows (Figure 7H).

Plants of this group are characteristically densely tuft-

ed with basally concentrated foliage, cauline leaves being

absent. The leaves are rolled or folded, narrow to fili-

form (1—3 mm wide), and glabrous. The inflorescences

are exserted well above the foliage. Bases are dense, com-

pact to almost flabellate with short rhizomes. Lateral

spreading rhizomes are seldom present and then only up

to 0,1 m long.

All voucher specimens were determined as C. dregeanus

and this group appears to represent a mountain form of

this species. A different concept of C. dregeanus appears

to have been applied by Chippindall (1955) as she also

included specimens with tall culms with cauline leaves

from the Namaqualand coastal plain. In the present study

this latter type of specimen has been grouped with C.

schraderi in Group 2. This nomenclatural problem must

be resolved before the correct names for these taxa can

be applied. Group 4 is nevertheless, morphologically,

ecologically and anatomically distinct and separate from

the other three groups.

CONCLUSIONS

The 33 Chaetobromus specimens examined in this

study can be grouped into four anatomical entities, each

possessing a characteristic combination of attributes;

most groups also have additional diagnostic anatomical

characters. Thus Group 1 has abaxial macrohairs and

tapering microhairs; Group 2 has truncated microhairs,

thick, flat or slightly infolded blades and distinct chloren-

chyma; Group 3 is similar to Group 1 but lacks macro-

hairs and microhairs and has sinuous long cell walls, and

Group 4 has setaceous leaves without second order vas-

cular bundles. Other characters are shared by pairs of

groups. Groups 1 and 2 both have microhairs and straight-

walled long cells whereas Groups 3 and 4 do not have

microhairs and the long cells have sinuous walls. Groups

1 and 3 both have inrolled leaves whereas Groups 2 and

4 tend to have infolded leaves.

Discontinuities are therefore not clear between the

four groups and they tend to intergrade into one another

in a reticulate manner. The groups are not discrete en-

tities but merely seem to reflect evolutionary trends. A

similar situation is also evident in the ecological prefer-

ences and vegetative and spike let morphology. For these

reasons it is extremely difficult to reliably and consistent-

ly define species and the recommendations of this study

are to uphold only a single species in the genus with four

reasonably clear subspecific taxa. These appear to re-

present ecotypes, each occurring in a specific habitat;

the differing ecological parameters appear to be reflect-

ed in the anatomy and vegetative morphology but no

consistent spikelet differences appear to be associated

with these environmental differences.

Populations of Chaetobromus plants in the field are

exceedingly variable, even within limited populations,

with each plant often differing in vegetative morphology.

This appears to be the case even in homogeneous habi-

tats where all the individuals appear to be of equal age

and are subject to the same environmental constraints.

In less uniform habitats, such as rocky outcrops where

soil depth, moisture availability and insolation can vary

over very short distances, this variation becomes even

more exaggerated, particularly if the area is also subject

to grazing. This type of variation suggests that genetic

differences exist between individuals even in small

populations. This appears to be confirmed by chromo-

some numbers of n = 12, 24 and 27 for different speci-

mens of Chaetobromus as well as indications of karyo-

type evolution in the form of paracentric inversions

(Spies & Du Plessis 1988).

Clearly the taxonomy of this genus requires further

biosystematic study including detailed fieldwork, cyto-

genetics and even transplant experiments. This detailed

study is fully justified as Chaetobromus appears to

possess excellent potential as a fodder grass. With correct

veld management based on knowledge of the reproduc-

tive biology and vegetative ontogeny of the genus this

grass could help considerably in enhancing the range

quality and carrying capacity of the Succulent Karoo

Biome.

ACK NOW LEDGEMENTS

Mrs H. Ebertsohn is gratefully thanked for technical

assistance, Mrs A. Romanowski for the photography,

Mrs S. Perold for the electron microscopy, and Mrs M.

van der Merwe for typing the manuscript.

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Bothalia 18,2: 211-220 (1988)

Flora of the Zuurberg National Park. 1. Characterization of major

vegetation units

B-E. VAN WYK*, P.A. NOVELLIE** and C.M. VAN WYK+

Keywords: Afromontane Forest, biogeography, eastern Cape, Fynbos Biome, Grassland, Grassy Fynbos, Mountain Fynbos, Subtropical

Thicket, vegetation units

ABSTRACT

The distribution of major vegetation units or veld types in the Zuurberg National Park, situated on the eastern

limits of the Fynbos Biome, is presented. Structural and floristic criteria are used to describe and map five basic

units, namely Afromontane Forest, Subtropical Thicket, Mountain Fynbos, Grassy Fynbos and Grassland.

UITTREKSEL

Die verspreiding van die hoofplantegroei-eenhede of veldtipes in die Zuurberg Nasionale Park, gelee aan die

oostelike grens van die Fynbosbioom, word aangebied. Strukturele en floristiese maatstawwe word gebruik om

vyf basiese eenhede te beskryf en te karteer, naamlik Afromontane Woud, Subtropiese Ruigtes, Bergfynbos,

Grasryke Fynbos en Grasveld.

INTRODUCTION

The Zuurberg National Park represents one of the

largest conservation areas incorporating Grassy Fynbos,

a vegetation type characteristic of the eastern limits of

the Fynbos Biome. An intricate mosaic of vegetation

types is present, reflecting the rugged topography, variety

of aspects and different microclimates. The biogeographi-

cal complexity of the eastern Cape is well known and is

a result of the convergence of four major phytochoria

(Goldblatt 1978; Gibbs Russell & Robinson 1981;

Cowling 1983a, 1983b, 1984; Lubke et al. 1986). This

diversity represents a major challenge in terms of con-

servation, since management measures taken for one

plant community may not be suitable for another.

* Department of Botany, Rand Afrikaans University, P.O. Box

524, Johannesburg 2000.

** National Parks Board, Private Bag X66, Cradock 5880.

t Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

MS. received: 1987.12.28.

No comprehensive account of the vegetation of the

Park is available. Existing knowledge is fragmentary and

limited to unpublished official reports and management

plans (Stehle 1979; Charlton 1982; Breytenbach &

Vlok 1985; Geldenhuys 1985). A research project to

provide basic floristic data and to explore the vegeta-

tional diversity was started in 1985. The results of a

preliminary survey aimed at describing and mapping the

major vegetation units is presented in this paper.

STUDY AREA

The study area is situated in the Zuurberg mountain

range, approximately 70 km due north of Port Eliza-

beth (Figure 1). The Park comprises three separate

parts with a total area of more than 20000 ha. The

Zuurberg forms part of the Cape Folded Belt and con-

sists mainly of hard quartzitic rock of the Witteberg

Group with numerous narrow bands of shale. Most of

the area is characterized by a series of mountain plateaus

separated by deep valleys with an east-west orientation.

FIGURE 1. — Locality of the stu-

dy area showing the dif-

ferent parts of the Zuurberg

National Park.

Both alia 18,2 (1988)

212

Bothalia 18,2 (1988)

213

TABLE 1. — Mean monthly rainfall (mm) at Zuurberg Lot 16 for the period 1931 to 1962 (Geldenhuys 1985)

JFMAMJ JASOND Total

66,8 63,8 95,0 53,3 43,0 25,5 38,3 35,7 77,0 79,7 78,9 64,7 721,7

The topography is very rugged due to the erosion of

softer shale bands from between alternating layers of

quartzites, but there are no peaks or steep cliffs. Height

above sea level varies between 250 and 970 m. The climate

is temperate with a mean annual rainfall of ± 722 mm.

Table 1 shows rainfall figures recorded at the office (Lot

16) between 1931 and 1962. The mean monthly figures

clearly show that spring and autumn maxima are expe-

rienced. Unlike other fynbos areas, winter months are

the driest. Thunderstorms commonly occur during the

summer months, when lightning fires may also be ex-

pected. Soil texture and soil depth vary considerably as

a result of the geological and topographical diversity.

The soils of the Zuurberg are generally more fertile and

finer textured than soils of the Cape Folded Belt to the

west (Campbell 1983; Cowling 1984).

METHODS

Ground patrols and an aerial reconnaissance by heli-

copter were undertaken to interpret aerial photographs

and to become familiar with the terrain, vegetation and

plant species. Herbarium specimens were collected on

several visits during 1985, 1986 and 1987. A checklist

of all the plant species recorded, complete with their

author names, is presented in part 2 of this series (Van

Wyk et al. 1988).

Vegetation units of the study area were visually iden-

tified according to vegetation structure and they corres-

pond roughly with the ‘veld type’ concept of Acocks

(1953) or the rank of class (Cowling 1984). A map

(Figure 2) was drawn from 1:50000 aerial photographs.

A series of colour slides taken by helicopter from differ-

ent angles at low altitude was used to verify the boun-

daries between vegetation units. The area covered by each

unit (Table 3) was estimated from the 1:50000 map by

a randomly positioned 2 mm grid. Descriptive data for

each of the five major vegetation units were obtained

from 64 sample quadrats distributed as shown in Table

2. All the quadrats were permanently marked by 1,2 m

iron fencing standards in each corner (5 x 10 m plots),

one at the centre of each short end (4 x 25 m plots) or

one at the centre (400 m2 circular plots). Localities were

selected so as to include most of the variation in each

vegetation type. Plot size varied between 50 and 400 m2

(Table 2).

The following information was recorded in each plot:

all identifiable species present, Braun-Blanquet cover

values for each species (r = < 1 % projected canopy cover;

1 = <5%; 2 = 6-25%; 3 = 26-50%; 4 = 51-75%;

5 = >75%), total projected canopy cover of all species,

height of different strata (grass layer, shrub layer, canopy

height and height of emergents) and, for Forest and

Thicket plots, also diameter at breast height of all indi-

vidual trees (if more than 100 mm). In view of the tremen-

dous variability of the vegetation, the sample size was

inadequate for a detailed phytosociological classification.

It does, however, provide sufficient information to cha-

racterize the major vegetation units. For descriptive pur-

poses, species were classed into growth forms as shown

in Tables 5-14. Dominant and characteristic species of

each vegetation unit were chosen as follows:

Characteristic (diagnostic) species: species with a

fidelity value of 80% or more.

Dominant species: species with a mean Braun-

Blanquet cover value of at least 0,80.

In these calculations, single occurrences (species pre-

sent in only one plot) were excluded. Forest and Thicket

plots were considered separately from the Mountain Fyn-

bos, Grassy Fynbos and Grassland plots. This seemed

reasonable as only a few species were common to both

subdivisions, and of these very few qualified as character-

istic or dominant.

RESULTS

The dominant vegetation types in the study area were

Grassy Fynbos (33%) and Subtropical Thicket (32%)

(Figure 2, Table 3). Grassland (18%), Afromontane

Forest (12%) and Mountain Fynbos (5%) had more

limited distributions.

A summary of floristic and structural characteristics

of the major vegetation units as recorded in 64 sample

plots is given in Table 4. Species richness (expressed as

species per m2 of plot area) varied between 0,13 (Forest)

to 0,92 (Mountain Fynbos). These values are dependent

on quadrat area, so that only the figures for Grassland,

Grassy Fynbos and Mountain Fynbos are directly com-

parable. The high figure for Thickets compared to

Forest agrees with previous findings that Afromontane

Forests are poorer in species than Thickets in the east-

ern Cape (Cowling 1983b). When distinct differences

in structure (Table 4) are considered in conjunction with

diagnostic and dominant species (Tables 5 to 14), each

of the major units is clearly distinguishable.

1 . Afromontane Forest

Forests comprising tall evergreen trees with canopy

heights of 10 to 14 m and emergents of up to 21 m occur

on south-facing slopes and in some valley bottoms.

Forest types on northern slopes and in alluvial valley

bottoms with canopy heights of 2—9 m and emergents

of up to 12 m are grouped with the next unit (Sub-

tropical Thicket). The distinction was not made on the

basis of structure only. We also used the almost total

absence of a herbaceous ground layer and the presence

of typical Afromontane species (White 1978) such as

Podocarpus falcatus and Diospyros whyteana. Despite a

strong Tongoland-Pondoland influence, there are pro-

nounced floristic differences between Afromontane For-

est and Subtropical Thicket in the eastern Cape (Cowling

1984). In the results of our survey, 24 tree species have

fidelity values of more than 80% (present in less than

214

Bothalia 18,2 (1988)

TABLE 2. — Number of sample quadrats. All were permanently marked to double as long term monitoring plots

TABLE 3. — Distribution of major vegetation units in different parts of the Zuurberg National Park. Areas were estimated from the

maps in Figure 2

TABLE 4. — Floristic and structural characteristics of the major vegetation units of the Zuurberg National Park

Bothalia 18,2 (1988)

20% of non-Forest plots) and 18 of these were recorded

only in Forest. Shrubs are rare, succulents are virtually

absent, and eight of the 10 fern species recorded occur

exclusively in Forest. Diagnostic and dominant species

are listed in Tables 5 and 6.

2. Subtropical Thicket

Thicket as defined here comprises a variable assem-

blage of communities dominated by thorny and/or suc-

culent shrubs. They occur on dry north-facing slopes

in higher parts and on all aspects in lower-lying south-

ern parts of the study area. Most of them comprise

what Lubke et al. (1986) describe as Valley Bushveld.

This term was used by Acocks (1953) but is no longer

useful because it incorporates too wide a range of types

(Cowling 1984). At least three basic types were includ-

ed in our sample:

TABLE 5. — Diagnostic species of Afromontane Forest grouped

by growth form. Single occurrences are excluded

* Sum of Braun-Blanquet cover estimates divided by presence; for

Forest and Thicket plots a cover of less than 5% was recorded as

0,50.

215

Kaffrarian Thicket

Closed, non-succulent shrubland to low forest com-

munities dominated by evergreen, sclerophyllous trees

and shrubs with a high cover of stem spines and vines

(Cowling 1984; Everard 1987). Campbell (1985) would

classify much of the taller thicket of this type as Eastern

Forest & Thicket.

Kaffrarian Succulent Thicket

This type occurs in dry areas and is characterized by a

high proportion of succulents, a great diversity in growth

form and a strong Karoo-Namib floristic influence (Cow-

ling 1984; Campbell 1985; Everard 1987). A variation

of this type, similar to Addo Bush and Sundays River

Scrub (Acocks 1953), occurs in southern parts such as

the northern slopes at Superbus (see Figure 2). The lat-

ter has a canopy height of no more than 2—3 m and is

dominated by Schotia afra, Putterlickia pyracantha, Phyl-

lanthus verrucosus and Euphorbia ledienii. Portulacaria

afra dominates in some parts, particularly on steep slopes

in the western parts of the Park.

Combretum caff rum- Acacia caffra Thicket

This type has a very limited distribution along river

beds. Since the dominant species ( Combretum caffrum

and Acacia caffra) are deciduous, it is not included in

the Subtropical Thicket concept and we group it here

provisionally. It shows similarity to the tropical thickets

found in the valleys of Natal and Transvaal.

Diagnostic and dominant species of Subtropical

Thicket are listed in Tables 7 and 8.

3. Mountain Fynbos

Mountain Fynbos has the highest species richness

(Table 4) and covers only an estimated 5% of the Park

TABLE 6.— Dominant species of Afromontane Forest grouped

by growth form. Single occurrences are excluded

216

TABLE 7. — Diagnostic species of Thicket communities grouped

by growth form. Single occurrences are excluded

* Invasive exotic

(Table 3). It is most common on wet southern slopes,

but also occurs on sandy soils in protected low-lying

areas. Mountain Fynbos as defined here is largely synony-

mous to the Mountain Fynbos of Taylor (1978), and

Kruger (1979) and the Mesotrophic Proteoid Fynbos of

Campbell (1985), but it also includes some communities

that may represent later serai stages of the next unit

(Grassy Fynbos). The Mountain Fynbos represented in

our sampling has a relatively low total grass cover and a

high proportion of C3 grasses ( Festuca , Pentaschistis

and Merxmuellera species) and Restionaceae. Mature

stands of Pro tea lorifolia and P. repens are very localiz-

ed and occur mostly in inaccessible areas. Campbell

Bothalia 18,2 (1988)

TABLE 8.- — Dominant species of Thicket communities grouped

by growth form

(1985) did not include mature fynbos of the Zuurberg

in his sample, presumably because it was thought to be

absent (Campbell op. cit. , page 7).

Widdringtonia nodiflora did not occur in any of our

fynbos plots, but it is highly characteristic of our con-

cept of Mountain Fynbos and should be added to the

lengthy list of diagnostic species in Table 9. A large

number of species are locally dominant. The list of

dominant species (Table 10) has therefore been limited

to those with a mean cover value of 1 ,00 or more and,

except for species with very high cover values, a pre-

sence of more than 50% (present in at least eight of the

15 Mountain Fynbos plots).

4. Grassy Fynbos

Grassy Fynbos covers the largest proportion of the

surface area of the Park (Table 3) and occurs on all

plateau tops and also on gentle southern and northern

slopes in higher-lying areas. Diagnostic and dominant

species of Grassy Fynbos are listed in Tables 1 1 and 12.

Campbell (1985) distinguished between three sub-

series of Grassy Fynbos, namely Dry, Mesic and Meso-

trophic. He classified the dominant vegetation of the

Zuurberg as Sundays Mesic Grassy Fynbos but also men-

tioned the lack of good differential characters. The pre-

sence of proteoids over 1 m tall, the less than 40% cover

of Ericaceae, the 10—50% cover of restioids and the

30—90% cover of grasses are used as differentiating fea-

tures by Campbell. Two types of his Mesotrophic sub-

Bothalia 18,2 (1988)

217

TABLE 9. — Diagnostic species of Mountain Fynbos communi- TABLE 10. — Dominant species of Mountain Fynbos communi-

ties grouped by growth form. Single occurrences are ex- ties grouped by growth form. Criteria for entry are specified

eluded in the text

series also occur in the Zuurberg, namely Mannetjiesberg

Mesotrophic Grassy Fynbos and Grahamstown Meso-

trophic Grassy Fynbos. In the study area, these two types

have a much more limited distribution than Sundays

Mesic Grassy Fynbos.

Our concept of Grassy Fynbos is much wider than

that of Campbell. We also include Suurberg Grassland

and much of Hankey Grassland, both of which approach

Acocks’s (1953) Dohne Sourveld (Campbell op. tit.).

Hankey Grassveld shows two extremes. A sourveld with

Tristachya leucothrix, Merxmuellera stricta and nume-

rous fynbos elements and a sweetveld with grasses such

218

TABLE 11. — Diagnostic species of Grassy Fynbos grouped by

growth form. Single occurrences are excluded

TABLE 12. — Dominant species of Grassy Fynbos grouped

by growth form

Bothalia 18,2 (1988)

as Themeda triandra and Heteropogon contortus with-

out fynbos elements (Campbell op. cit.). The sourveld

is here included under Grassy Fynbos and the sweetveld

under Grassland. In the study area, there is a much great-

er discontinuity in the distribution of fynbos elements

than in those characters used by Campbell to distinguish

between Grassy Fynbos and Grassland. For practical

reasons, we have therefore used the presence of fynbos

elements to differentiate between Grassy Fynbos and

Grassland.

5. Grassland

R.A. Lubke (unpublished data) recognized, in the

northern part of the Sundays River area (Figure 1), two

major grassland communities, namely Festuca costata

Tussock Grassland and Themeda triandra- Tristachy a

leucothrix Grassland. The latter was provisionally sub-

divided by him into Bobartia orientalis Grassland and

Trachypogon spicatus Grassland, further subdivided into

a, Heteropogon contortus Grassland and b, Setaria spha-

celata Grassland.

We have taken a much narrower view and the Grass-

lands of the study area are here considered to include

only those areas where Restionaceae, Ericaceae and Pro-

teaceae are totally absent. Campbell’s (1985) criteria

for recognizing Grassland are difficult to use because of

the gradual decrease of fynbos elements along the tran-

sition from Grassy Fynbos to Grassland. Our concept

therefore includes only part of Campbell’s Hankey

Grassland and seems to be identical to Lubke’s Setaria

sphacelata Grassland. As such it is perhaps the most uni-

form vegetation unit of all and occurs mostly on steep

northern and western slopes. What variation there is,

appears to be the result of soil depth and rockiness.

Some species ( Acacia karroo, Diospyros lycioides and

Aloe ferox for example) are restricted to deep soils on

lower northern slopes, while succulents such as Euphor-

bia polygona are locally dominant only in very rocky

areas.

Diagnostic and dominant species are listed in Tables

13 and 14. Very few of the dominant species have high

fidelity values, so that most of the diagnostic species are

forbs or succulents and not grasses. Elionurus muticus

and Brachiaria serrata are very common but only Setaria

sphacelata var. torta and Aristida diffusa subsp. burkei

appear to be characteristic of Grassland as defined here.

The dominant grasses are also present in Grassy Fynbos,

where their cover values are scarcely lower.

DISCUSSION

The forests of the Zuurberg have floristic elements in

common with both the Amatola and Alexandria Forests

and are similar in species composition to forests in the

Watersmeeting Nature Reserve (Bathurst), the Fort Grey

Nature Reserve (East London) and the Groendal Wilder-

ness Area north of Uitenhage (Geldenhuys 1985). They

differ from the Knysna and Tsitsikamma Forests in species

composition, notably the absence of Ocotea bullata and

Trichocladus crinitus and the presence of species of Pon-

doland-Tongaland affinity. Trees such as Smellophyllum

capense, Atalaya capensis, Homalium dentatum, H. ru-

fescens and Chionanthus peglerae are rare or have limited

Bothalia 18,2 (1988)

TABLE 13. — Diagnostic species of Grassland grouped by growth

form. Single occurrences are excluded

distributions in the eastern Cape. The Zuurberg Forests

are relatively isolated and therefore represent an impor-

tant biogeographical link in the distribution of Afro-

montane Forest in the eastern Cape and also between

the eastern and southern Cape (Geldenhuys 1985).

Thicket communities of the Zuurberg are very vari-

able, probably as a result of topographic, rainfall and

edaphic gradients. In terms of structure and species

composition, most of the thicket agrees with Everard’s

(1987) Xeric Kaffrarian Thicket. Only a very small part

of the Kaffrarian Succulent Thicket of the study area is

similar to Addo Bush (Acocks 1953) or Spekboomveld

(Archibald 1955), the dominant vegetation of the Addo

Elephant National Park. Everard (1987) classified the

latter as one of two suborders of Kaffrarian Succulent

Thicket, namely Xeric Succulent Thicket. His other

suborder, Mesic Succulent Thicket, seems floristically

similar to some of the thickets of the Zuurberg.

The Mountain Fynbos of the study area was not in-

cluded in the classification of Campbell (1985) but it

has the differentiating features of his Mesotrophic Pro-

teoid Fynbos. Protea lorifolia and P. repens are the

dominant canopy species and Grassy Fynbos is present

as understorey. In view of the limited and localized dis-

tribution and the high species richness, this vegetation

unit should receive special attention when management

219

TABLE 14. — Dominant species of Grassland grouped by growth

form

policies are formulated. The effects of fire on succession

(under local conditions) need to be studied in detail.

The absence of seed-regenerating Protea species (P. lori-

folia and P. repens ) from the plateau areas may be part-

ly due to frequent fires in the past (see the successional

model of Cowling 1984), but shallower and more fertile

soils could also be limiting factors. Mature stands of

Protea lorifolia are almost exclusively found on sandy

soils.

Judged by diagnostic species, our concept of Grassy

Fynbos appears to be similar to that of Cowling (1984),

who argued that it is not a recently derived vegetation

type as Acocks (1953) proposed. The presence of regional

endemics {Erica demissa, E. pectinifolia, Podalyria bur-

chellii and Protea foliosa for example) and the resprout-

ing ability of virtually all the species found in Grassy

Fynbos indicate that it should be recognized as a distinct

vegetation type. Campbell (1985) proposed that the

Grassy Fynbos (Eastern Fynbos) of the Zuurberg and

Grahamstown areas should be included in the Fynbos

Biome and perhaps also in the Cape Floristic Region.

The abundance of grasses was discussed by Cowling

(1984). He suggested that high temperatures during the

growing season (the high proportion of summer rain) in-

crease the competitive advantage of C4 grasses, although

the more fertile and finer-textured soils (Campbell 1983)

also need to be considered. Too frequent fires may lead

to an increase in grassiness by removing the shading effect

of the overstorey. It is possible that longer intervals be-

tween fires will result in an increase of Mountain Fynbos

in certain areas.

220

CONCLUSIONS

In the Zuurberg National Park several totally differ-

ent and unrelated vegetation types occur in close proxi-

mity. The dynamics of the boundaries between the types

and between communities need to be studied in more

detail to explain the intricate mosaic of vegetation. Com-

munity boundaries may be determined at least partly by

an equally intricate mosaic of soil types. Campbell (1983)

has reported distinct edaphic gradients in the mountains

of the Fynbos Biome. Another major ecological factor

seems to be the natural fire cycle, in which the warmer,

drier northern slopes tend to burn at more frequent

intervals than the wet southern slopes. Fire is considered

to be the major disturbance factor in fynbos biome com-

munities (Cowling et al. 1987).

If the present-day patchy distribution of plant com-

munities has been shaped by edaphic factors and a natural

fire regime, little seems to be gained by interfering with

the natural cycle. It is indeed impractical to divide such

complex communities into conventional ‘burning blocks’

and burn them according to a rigid schedule. Instead,

management should try to allow lightning fires to run

their natural course, and exclude man-made fires from

outside the Park. Further research is required to assess

this policy, focusing on post-fire succession and the

effects of fire on the characteristic and dominant species

in each vegetation unit. It is also important to gain infor-

mation on the extent to which edaphic factors are re-

sponsible for community boundaries.

ACKNOWLEDGEMENTS

We thank Nico and Sorina van der Walt for their hos-

pitality during our visits. Financial support from the

National Parks Board and the Rand Afrikaans University

is acknowledged.

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inheemse woudplantegroei in the Suurberg Nasionale Park.

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graphy and speciation in the vegetation of the eastern

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VAN WYK, B-E., VAN WYK, C.M. & NOVELLIE, P.A. 1988.

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Bothalia 18,2: 221-232 (1988)

Flora of the Zuurberg National Park. 2. An annotated checklist of ferns

and seed plants

B-E. VAN WYK*, C.M. VAN WYK** and P.A. NOVELLIEt

Keywords: checklist, eastern Cape, flora, growth form, plant distribution

ABSTRACT

A checklist of 1 100 plant taxa representing 540 genera and 129 families is given for the Zuurberg National Park

(eastern Cape). Annotations include growth form (except in Poaceae and Cyperaceae) and voucher specimen(s).

The diversity of the flora is ascribed to the presence of four major phytochoria of southern Africa rather than

local endemism.

UITTREKSEL

’n Kontrolelys van 1 100 planttaksons, wat 540 genusse en 129 families verteenwoordig, word vir die Zuur-

berg Nasionale Park (Oos-Kaap) aangegee. Groeivorms (behalwe by Poaceae en Cyperaceae) en kontrole-eksemplare

word ook aangetoon. Die diversiteit van die flora word toegeskryf aan die teenwoordigheid van vier hoofplante-

groeistreke van suidelike Afrika eerder as aan plaaslike endemisme.

INTRODUCTION

The study area is situated in the eastern Cape, which

is known to be a transitional zone of four major phyto-

choria, namely the Cape, Tongoland-Pondoland, Karoo-

Namib and Afromontane regions (Goldblatt 1978; Gibbs

Russell & Robinson 1981; Cowling 1983; Lubkeefa/.

1986). Furthermore, five vegetation types are represent-

ed in this area. It is therefore of considerable phytogeo-

graphical interest.

The vegetation of the Zuurberg National Park was

studied to supply basic data for an inventory. Major

vegetation units are described by Van Wyk etal. (1988)

and all ferns and flowering plants recorded within the

boundaries of the Park are listed. No comprehensive

studies or systematic collecting work have been done

here in the past.

METHODS

Herbarium specimens were collected on several visits

between June 1985 and October 1987. The staff of the

National Herbarium and various specialists identified the

material. A field herbarium was compiled to provide a

permanent record of all plant taxa and to facilitate future

studies. Duplicates of the specimens are housed in the

Rand Afrikaans University Herbarium (JRAU) and in

the National Herbarium (PRE) of the Botanical Research

Institute, Pretoria.

The nomenclature and arrangement of the checklist

follow Gibbs Russell et al. (1985, 1987). Collecting

* Department of Botany, Rand Afrikaans University, P.O. Box

524, Johannesburg 2000.

** Botanical Research Institute, Department of Agriculture and

Water Supply, Private Bag X101, Pretoria 0001.

t National Parks Board, Private Bag X66, Cradock 5880, South

Africa.

MS. received: 1987.12.28.

numbers, all by B-E. & C.M. van Wyk, are shown in

italics. Growth forms are mentioned, except in Poaceae

and Cyperaceae. Naturalized exotic species are marked

with an asterisk. Exact localities are not cited, but all

collections are from the following quarter degree square

grids: 3325 (Port Elizabeth): —AD, — BC, — BD.

RESULTS

A total of 1 100 species and infraspecific taxa repre-

senting 540 genera and 129 families, was recorded in the

study area (Table 1). Naturalized exotic species represent

5,6% of the total flora, a figure much lower than the

10% given by Gibbs Russell & Robinson (1981) for the

Albany and Bathurst Districts. The largest families are

the Asteraceae (62 genera, 148 species) and the Poaceae

(51 genera, 90 species). Nearly all of the largest families

of southern Africa (Gibbs Russell 1985) are also present

amongst the 50 largest families of the Zuurberg. Only

the Mesembryanthemaceae and the Ericaceae are ranked

significantly lower in the Zuurberg flora than in the

southern African flora as a whole (Gibbs Russell 1985,

Table 3).

TABLE 1. — Number of families, genera and species recorded in

the Zuurberg National Park

222 Bothalia 18,2 (1988)

TABLE 2. — Comparison of the flora of the Zuurberg National Park with other southern African floras (data from Gibbs Russell 1985)

The diversity of the flora is compared to other south-

ern African floras in Table 2. The Zuurberg National

Park is much smaller than the other areas mentioned in

Table 2. However, more vegetation types are represented

than in the other regional floras mentioned (Van Wyk et

al. 1988) and a comparable number of families and genera

is present. Although the number of species per km2 is

very high, the total number of species and the species/

genus ratio are lower. Endemic genera typically have

high species/ genus ratios. A mean of 8,6 for all endemic

genera in southern Africa is given by Gibbs Russell &

Robinson (1981). A lack of distribution records from

many of the quarter degree grid squares of the eastern

Cape makes it very difficult to accurately assess the

degree of endemism. Lubke et al. (1986) have concluded

that endemism in the eastern Cape is relatively low com-

pared to other centres. Their results have indicated that

most endemics (61) are from the thicket vegetation of

the Tongaland-Pondoland region, with the Cape region

(59) a close second. The diversity of the flora of the

Zuurberg National Park is ascribed not so much to

endemism or to a large number of species, but rather to

the presence of five vegetation types and the relatively

high number of higher order taxa (families and genera),

each of which contribute a relatively low number of

species.

ACKNOWLEDGEMENTS

We wish to thank the Director and staff of the National

Herbarium for the many specimens that were identified.

The following specialists have also contributed to iden-

tifications: A. Bean, K. Balkwill, H.P. Linder, R.O. Mof-

fett, A.A. Obermeyer-Mauve, E.G.H. Oliver, C.H. Stirton

and A.E. van Wyk. We also wish to thank the following

persons who helped us in various ways: H. Allison, R.M.

Cowling, M. Fourie, L. Jonker, M.Koekemoer, G.Kotze,

R.A. Lubke, R. Niemand, E. Novellie, E.M. Reinten, A.L.

Schutte, N. van der Walt, S. van der Walt and K.T. van

Warmelo. Financial support from the National Parks

Board and the Rand Afrikaans University is acknow-

ledged.

REFERENCES

COWLING, R.M. 1983. Phytochorology and vegetation history

in the south-eastern Cape, South Africa. Journal of Bio-

geography 10: 393-419.

GIBBS RUSSELL, G.E. 1985. Analysis of the size and composi-

tion of the southern African flora. Bothalia 15: 613 —

629.

GIBBS RUSSELL, G.E. & ROBINSON, E.R. 1981. Phytogeogra-

phy and speciation in the vegetation of the eastern Cape.

Bothalia 13: 467-472.

GIBBS RUSSELL, G.E., REID, C., VAN ROOY, J. & SMOOK,

L. 1985. List of species of southern African plants, edn 2,

part 1. Memoirs of the Botanical Survey of South Africa

No. 51.

GIBBS RUSSELL, G.E., WELMAN, W.G., RETIEF, E., IMMEL-

MAN, K.L., GERMISHUIZEN, G., PIENAAR, B.J., VAN

WYK, M. & NICHOLAS, A. 1987. List of species of south-

ern African plants, edn 2, part 2. Memoirs of the Botanical

Survey of South Africa No. 56.

GOLDBLATT, P. 1978. An analysis of the flora of southern

Africa: its characteristics, relationships and origins. Annals

of the Missouri Botanical Garden 65: 369-436.

LUBKE, R.A., EVERARD, D.A. & JACKSON, S. 1986. The

biomes of the eastern Cape with emphasis on their conser-

vation. Bothalia 16: 251-261.

VAN WYK, B-E., NOVELLIE, P.A. & VAN WYK, C.M. 1988.

Flora of the Zuurberg National Park. 1. Characterization

of major vegetation units. Bothalia 18: 211—220.

CHECKLIST

The nomenclature and arrangement of the checklist follow Gibbs Russell et al. (1985, 1987). Collecting numbers, all by B. & M.

van Wyk, are shown in italics. Growth forms are mentioned, except in ferns, Poaceae and Cyperaceae. Naturalized exotic species are

marked with an asterisk. Exact localities are not cited, but all collections are from the following quarter degree square grids: 3325

(Port Elizabeth): -AD, -BC, -BD.

PTERIDOPHYTA

EQUISETACEAE

Equisetum ramosissimum Desf, 1963

OSMUNDACEAE

Todea barbara (L.) T. Moore, 21 73

SCHIZAEACEAE

Mohria caffrorum (L.) Desv., 231, 340

Schizaea pectinata (L.) Swartz, 183, 1283, 1517

DENNSTAEDTIACEAE

Pteridium aquilinum (L.J Kuhn subsp. aquilinum, 228a

ADLANTACEAE

Adiantum

capillus-veneris L., 326

raddianum Presl, 331

Cheilanthes

capensis (Thunb.) Swartz, 1378

concolor (Langsd & Fisch.) Schelpe & N.C. Anthony, 777,

1088

hirta Swartz, 1768, 1886

multifida (Swartz) Swartz var. multifida, 1238

parviloba (Swartz) Swartz, 482, 1086

viridis (Forssk.) Swartz

var. macrophylla (Kuntze) Schelpe & N.C. Anthony, 334

var. viridis, 369, 642, 695, 1083, 1087

Bothalia 18,2 (1988)

223

Pellaea calomelanos (Swartz) Link, 703, 964

Pteris dentata Forssk., 337

POLYPODIACEAE

Pleopeltis macrocarpa (Bory ex Willd) Kaulf, 749, 1084

ASPLENIACEAE

Asplenium

lunulatum Swartz, 349, 795, 1655

rutifolium (Berg.) Kunze, 1 77

splendens Kunze, 350

Ceterach cordatum (Thunb.) Desv., 339, 1356

THELYPTERIDACEAE

Thelypteris bergiana (Schlechtd) Ching, 769

LOMARIOPSIDACEAE

Elaphoglossum acrostichoides (Hook. <6 Grev.) Schelpe, 1 085

ASPIDIACEAE

Dryopteris inaequalis (Schlechtd.) Kuntze, 347

Rumohia adiantiformis (G. Forst.) Ching, 168

BLECHNACEAE

Blechnum

australe L. var. australe, 766, 1240

capense (L.) Schlechtd., 765, 2171

punctulatum Swartz var. punctulatum, 352, 1239

GYMNOSPERMAE

ZAMIACEAE

Encephalartos

caffer (Thunb.) Lehm., shrub, 2161

lehmannii Lehm., shrub, 626

longifolius (Jacq.) Lehm., small tree, 233

PODOCARPACEAE

Podocarpus

falcatus (Thunb.) R. Br. ex Mirb., tree, 753

latifolius (Thunb.) R. Br. ex Mirb., tree, 82

CUPRESSACEAE

Widdringtonia nodiflora (L.) Powrie, shrub or small tree, 200

ANGIOSPERMAE — MONOCOTYLEDONEAE

TYPHACEAE

Typha capensis (Rohrb.) N.E. Br., forb, 1629

POTAMOGETONACEAE

Potamogeton thunbergii Cham. & Schlechtd., aquatic forb, 238

APONOGETONACEAE

Aponogeton desertorum Zeyh. ex Spreng. f, aquatic forb, 1 985

POACEAE

Hemarthria altissima (Poir.) Stapf & C.E. Hubb., 1984

Elionurus muticus (Spreng.) Kunth, 1095b, 1120, 1287

Miscanthus

erectus (Stent & C.E. Hubb.) ined, 1480

sorghum (Nees) Pilg., 1584

Andropogon appendiculatus Nees, 1054, 1277

Cymbopogon

plurinodis (Stapf) Stapf ex Burtt Davy, 86 7

validus (Stapf) Stapf ex Burtt Davy, 707, 1196, 1200, 1318

Hyparrhenia

anamesa Clayton, 639, 1197

hirta (L.) Stapf, 1141

Trachypogon spicatus (L. f.) Kuntze, 1100

Heteropogon contortus (L.) Roem. & Schult., 669, 999, 1099

Diheteropogon filifolius (Nees) Clayton, 656, 1105, 1199

Themeda triandra Forssk., 6 73

Digitaria

eriantha Steud., 997, 1320, 1392

monodactyla (Nees) Stapf, 974, 1014, 1095a, 1424

natalensis Stent, 1126

Alloteropsis semialata (R. Br.) Hitchc. subsp. eckloniana (Nees)

Gibbs Russell, 611, 983, 1108

Brachiaria

arrecta (Dur. & Schinz) Stent, 1587b

serrata (Thunb.) Stapf, 995, 1143, 1340

Paspalum

dilatatum Poir., 1046, 1109*

paspaloides (Michx.) Scribm, 1587a

scrobiculatum L., 1043

Urochloa panicoides Beauv., 1764, 1941

Echinochloa crus-galli (L.) Beauv., 1955

Oplismenus sp., 1090, 1898

Panicum

coloratum L. var. coloratum, 1 741

deustum Thunb., 760, 978, 1398

ecklonii Nees, 1717

gilvum Launert, 1042, 1586

maximum Jacq., 963, 989, 1142, 1360, 1548

subalbidum Kunth, 2033

Setaria

lindenbergiana (Nees) Stapf, 891, 1323

sphacelata (Schumach.) Moss

var. sphacelata, 1366

var. torta (Stapf) Clayton, 965, 1198

Rhynchelytrum nerviglume (French.) Chiov., 218, 675, 969, 1503

Pennisetum macrourum Trim , 1220, 1988

Cenchrus ciliaris L., 1821

Ehrharta

calycina J.E. Sm. var. calycina, 1185, 1202b

erecta Lam. var. erecta, 1203, 1268

ramosa (Thunb.) Thunb. var. ramosa, 1223, 1276

cf. longifoha Schrad, 1274

Phalaris

arundinacea L., 1978*

minor Retz, 1918*

Arundinella nepalensis Trim, 2148

Tristachya leucothrix Nees, 66 7, 998

Helictotrichon hirtulum (Steud.) Schweick., 1053, 1102, 1265

Merxmuellera

arundinacea (Berg) Conert, 1836

disticha (Nees) Conert, 2101

stricta (Schrad) Conert, 865, 2053, 2102

Karroochloa curva (Nees) Conert £ Turpe, 135 7

Pentaschistis

ampla (Nees) McClean, 1060

angustifolia (Nees) Stapf, 1028, 1101, 1155, 1186, 1254, 1273

curvifolia (Schrad) Stapf, 1263, 1264

eriostoma (Nees) Stapf, 1316, 1837

Phragmites australis (Cav.) Steud, 1989

Agrostis lachnantha Nees var. lachnantha, 763a, 889

Polypogon monspeliensis (L.) Desf, 888*

Aristida

congesta Roem. & Schult. subsp. barbicollis (Trim & Rupr.) De

Winter, 1847

diffusa Trim subsp. burkei (Stapf) Meld., 863, 987, 1328

junciformis Trim & Rupr. subsp. junciformis, 1510

Stipa dregeana Steud var. elongata (Nees) Stapf 96 7, 1149, 1408,

1652

Tragus berteronianus Schult., 1363

Sporobolus

africanus (Poir.) Robyns & Toumay, 1050

centrifugus (Trim) Nees, 2108

fimbriatus (Trim) Nees var. fimbriatus, 1153, 1359

mauritianus (Steud.) Dur. & Schinz, 2068, 2105

natalensis (Steud) Dur. <6 Schinz, 1029

nitens Stent, 1386

Eragrostis

capensis (Thunb.) Trim, 471, 709, 742, 1271

cilianensis (AIL) F.T. Hubb., 1399, 1416

curvula (Schrad) Nees, 980, 1032, 1148, 1746

obtusa Munro ex FicaL & Hiem, 1093, 1145, 1391

plana Nees, 1065

planiculmis Nees, 242, 1052, 1057, 1218, 1221

racemosa (Thunb.) Steud, 281, 1037, 1098

Microchloa caffra Nees, 1269

Cynodon dactylon (L.) Pers., 1048, 1106, 1107, 1255

Harpochloa falx (L. f.) Kuntze, 1187

Eleusine coracana (L.) Gaertm subsp. africana (K.-OByme) Hilu

& De Wet, 1942

Triraphis andropogonoides (Steud.) PhilL, 131 7

Koeleria capensis (Steud) Nees, 1272, 1970, 1980

Melica racemosa Thunb., 994, 1202a, 1545

Schismus barbatus (Loefl ex L.) ThelL, 923

224

Bothalia 18,2 (1988)

Poa binata Nees, 2100

Festuca

costata Nees var. costata, 2098

scabra Vahl, 1665, 1723

Bromus unioloides H. B. K„ 1266, 1904, 1959*

Brachypodium flexum Nees, 262

Lolium rigidum Gaudin, 1 950*

CYPERACEAE

Carpha glomerata (Thunb.) Nees, 1233

Cyperus

albostriatus Schrad., 230, 972, 1939

denudatus L. f, 1044, 1585

difformis L., 1396

maiginatus Thunb., 929, 1806

obtusiflorus Vahl vai. sphaerocephalus (Vahl) Kuekenth., 1056

pulcher Thunb., 1 777, 1 778

rubicundus Vahl, 1393

sphaerosperm us Schrad., 1946

tenellus L. f. var. tenellus, 203 7b

textilis Thunb., 322

usitatus Burch., 1394

Pycreus polystachyos (Rottb.) Beauv., 1613

Mariscus

dregeanus Kunth, 1138

indecorus (Kunth) Podlech, 990

solidus (Kunth) ined.

vai. solidus, 1 792

var. involutus (C.B. Cl) ined., 1121

tabularis (Schrad.) C.B. Cl subsp. major (Nees) ined., 1397,

1770

uitenhagensis Steud., 977

usitatus (Burch.) ined. var usitatus, 1394

Ficinia

albicans Nees, 1827

quinquangularis Boeck., 1846

sylvatica Kunth, 1889

Fuirena hirsuta (Berg.) P.L. Forbes, 1579, 1956, 2143

Schoenoplectus paludicola (Kunth) Palla ex J. Raynal, 232, 2031

Isolepis

cemua (Vahl) Roem. <£ Schult., 896

ludwigii Kunth, 1673

prolifer R. Br., 783

sepulcralis Steud., 214, 881, 2037a

Eleocharis dregeana Steud, 2032

Fimbristylis complanata (Retz.) Link, 1047

Bulbostylis

contexta (Nees) Bodard, 1000, 1097

humilis (Kunth) C.B. CL, 1003

Abildgaardia ovata (Burm. f.) Krai., Femiebrae plot A 1

Tetraria

bromoides (Lam.) Pfeiffer, 2158

compressa Turrill, 264

cuspidata (Rottb.) C.B. CL, 357, 677, 744

fourcadei Turrill & Schonl., 2058

pubescens Schonl. & Turrill, 151 9

secans C.B. CL, 1520

sp. nov., 1499

Schoenoxiphium

lehmannii (Nees) Steud., 1735

rufum Nees, 21 79

sparteum (Wahlenb.) C.B. CL, 1049, 1506, 1471

Carex

cf. aethiopica Schkuhr, 1631

clavata Thunb., 1979

ARACEAE

Zantedeschia aethiopica (L.) Spreng., forb, 752a

LEMNACEAE

Lemna gibba L., aquatic forb, 934a

Wolffia arrhiza (L.) Horkel ex Wimm., aquatic forb, 934b

RESTIONACEAE

Ischyrolepis sieberi (Kunth) Linder, 2059, 2060, 2167

Elegia vaginulata Mast., 1521

Restio

sejunctus Mast., 1515

triticeus Rothb., 9, 267, 359

Calopsis paniculata (Rottb.) Desv., 2157

Rhodocoma

capensis Nees ex Steud., 125, 179, 1338, 1727, 2047

fruticosa (Thunb.) Linder, 420, 1 728

Cannomois virgata (Rottb.) Steud, 2046

Hypodiscus

striatus (Kunth) Mast., 268, 762, 768, 1514

synchroolepis (Steud.) Mast., 236

sp., 2182

COMMELINACEAE

Commelina

africana L.

var. africana, forb, 1658

var. krebsiana (Kunth) C.B. CL, forb, 944

var. lancispatha C.B. CL, forb, 946, 1390

benghalensis L„ forb, 948

eckloniana Kunth, forb, 1079, 1384

Cyanotis speciosa (L. f.) Hassk., forb, 992, 1286, 1379

JUNCACEAE

Prionium serratum (L. f.) Drkge ex E. Mey., large tuft, 2142

Juncus

capensis Thunb., tuft, 937

exsertus Buchen., tuft, 892

lomatophyllus Spreng., tuft, 1628

oxycarpus E. Mey. ex Kunth, tuft, 1 765, 2034

LILIACEAE

Bulbine

abyssinica A. Rich., succulent, 1068

asphodeloides (L.) Willd, succulent, 1677

filifolia Bak., forb, 1176

frutescens (L.) Willd, succulent, 323, 2150

cf. lagopus (Thunb.) N.E. Br., succulent, 2111

latifolia (L. f.) Roem. & Schult., succulent, 134

Trachyandra

affinis Kunth, forb, 21 77

asperata Kunth.

var. macowanii (Bak.) Oberm., forb, 1661

var. stenophylla (Bak.) Oberm., forb, 1055

gif fenii (Leighton) Oberm., forb, 1795

saltii (Bak.) Oberm. var. saltii, forb, 1707

Anthericum sp., forb, 21 77

Chlorophytum

capense (L.) Koss, forb, 1122

comosum (Thunb.) Jacq., forb, 1760, 1897

Caesia contorta (L. f.) Dur. <6 Schinz, forb, 1013

Schizobasis intricata (Bak.) Bak., geophyte, 1313, 1541

Eriospermum

dyeri Mar loth ex Archib., geophyte, 1312

sp. cf. E. occultum Archib., geophyte, 1475, 1022b, 2125

Kniphofia cf. uvaria (L.) Hook., forb, 370, 1429

Aloe

ferox Mill, large succulent, 156

microcantha Haw., succulent, 1224, 2156

pluridens Haw., large succulent, 1644

speciosa Bak., large succulent, 710

striata Haw., succulent, 621

tenuior Haw., scandent succulent, 40, 405

Gasteria sp., succulent, 2023

Haworthia

angustifolia Haw., succulent, 693, 1249, 1311

cooperi Bak. var. cooperi, succulent, 623, 734

glauca Bak. var. glauca, succulent, 732

reinwardtii (Salm-Dyck) Haw.

var. reinwardtii, succulent, 485

cf. var. brevicula G. G. Sm. , succulent, 1546

Agapanthus praecox Willd. subsp. minimus (Lindl.) Leighton, forb,

637, 1458

Tulbaghia

cemua Ave-Lall., geophyte, 1127, 1726

violacea Harv., forb, 1531

Albuca

sp. cf. A. macowanii Bak., geophyte, 691, 2151

sp., geophyte, 2107

Urginea altissima (L. f.) Bak., geophyte, 1423h, 1742

Drimia

anomala (Bak.) Benth., geophyte, 1843

capensis (Burm. f.) Wijnands, geophyte, 1315

sp., geophyte, 2011

Bothalia 18,2 (1988)

225

Rhadamanthus sp. nov.?, geophyte, 1858

Dipcadi viride (L.) Moench, geophyte, 722, 2003

Litanthus pusillus Harv., geophyte, 1571

Scilla nervosa (Burch.) Jessop, geophyte, 1010

Eucomis autumnalis (Mill) Chitt. subsp. autumnalis, geophyte,

2126

Omithogalum

anguinum Leighton ex Oberm., geophyte, 1653

conicum Jacq. subsp. conicum, geophyte, 1 722

juncifolium Jacq., geophyte, 1022a, 1188, 1842

rogersii Bak. , geophyte, 1890

Ledebouria

revoluta (L. f.) Jessop, geophyte, 1690

undulata (Jacq.) Jessop, geophyte, 692

Veltheimia bracteata Harv. ex Bak., geophyte, 1801, 2202

Sansevieria hyacinthoides (L.) Druce, forb, 630

Protasparagus

aethiopicus (L.) Oberm., climber, 536, 1636

burchellii (Bak.) Oberm., forb, 561

capensis (L.) Oberm., forb, 406

crassicladus (Jessop) Oberm., climber, 578, 806

densiflorus (Kunth) Oberm., forb, 521, 6 76

divaricatus Oberm., forb, 415

laricinus (Burch.) Oberm., forb, 646

macowanii (Bak.) Oberm., forb, 408, 1593

multiflorus (Bak.) Oberm., climber, 1614

setaceus (Kunth) Oberm., climber, 112, 341

striatus (L. f.) Oberm., forb, 116, 235, 519

suaveolens (Burch. ) Oberm., forb, 88, 364

subulatus (Thunb.) Oberm., forb, 555

virgatus (Bak.) Oberm., forb, 18, 343

Behnia reticulata (Thunb.) Didr., forb, 69

HAEMODORACEAE

Lanark lanata (L.) Dur. & Schinz, forb, 1034

AMARYLLIDACEAE

Haemanthus albiflos Jacq. , geophyte, 162, 1575

Scadoxus puniceus (L.) Friis & Nordal, geophyte, 1229

Boophane disticha (L. f.) Herb., geophyte, 694

Nerine undulata (L.) Herb, complex, geophyte, 1625

Brunsvigia grandiflora LindL, geophyte, 1470

Cyrtanthus sanguineus (LindL) Walp., geophyte, 2016

HYPOXIDACEAE

Empodium sp., geophyte, 1489

Hypoxis

angustifolia Lam. var. buchananii Bak., geophyte, 1944

hemerocallidea Fisch. & Mey., geophyte, 1008

longifolia Bak., geophyte, 1250, 1257

obtusa Burch, ex Edwards, geophyte, 862a

rigidula Bak. var. rigidula, geophyte, 1036, 2039

cf. villosa L. f., geophyte, 862

zeyheri Bak., geophyte, 1260

TECOPHILAEACEAE

Cyanella sp., geophyte, 802

DIOSCOREACEAE

Dioscorea cf. sylvatica (Kunth) Eckl, geophyte, 478

IRIDACEAE

Romulea autumnalis L. BoL, geophyte, 219

Moraea

cf. algoensis GoldbL, geophyte, 2043

cf. elliotii Bak., geophyte, 1710

tripetala (L. f.) Ker-GawL, geophyte, 2180

Dietes iridioides (L.) Sweet ex Klatt, forb, 1761

Bobartia orientalis J.B. Gillett subsp. orientalis, forb, 379, 2113

Aristea

anceps Eckl. ex Klatt, forb, 1020, 1245

cognata N.E. Br. ex Weim., forb, 1247

ecklonii Bak., forb, 1683

juncifolia Bak. , forb, 1509

schizolaena Harv., forb, 1123, 1259, 1696

Dierama pendulum Bak., geophyte, 17, 2138

Gladiolus ochroleucus Bak. var. macowanii (Bak.) Oberm., geo-

phyte, 1428

Anapalina caffra (Ker-GawL ex Bak.) G.J. Lewis, geophyte, 2

Watsonia cf. longifolia Mathews & L. BoL, geophyte, 658, 713

Freesia corymbosa (Burm. f.) N.E. Br., geophyte, 71 7

ORCHIDACEAE

Habenaria cf. arenaria LindL, forb, 1596, 1611

Brachycorythis macowaniana Reichb. f, forb, 1998

Disa versicolor Reichb. f. , forb, 2203

Satyrium membranaceum Swartz, forb, 1 720

Monadenia brevicomis LindL, forb, 1 706

Disperis lindleyana Reichb. f, forb, 1598

Acrolophia capensis (Berg.) Fourc. var. capensis, forb, 1691, 1999

Polystachya pubescens Reichb. f, epiphyte, 1163

Eulophia

clavicomis LindL var. clavicomis, forb, 65 7, 789

platypetala LindL, forb, 662

tenella Reichb. f, forb, 1177, 1190

Tridactyle tricuspis (H. BoL) Schltr., epiphyte, 1590, 2014

Cyrtorchis arcuata (LindL) Schltr., epiphyte, 2013

Mystacidium

capense (L. f.) Schltr., epiphyte, 2012

venosum Harv. ex Rolfe, epiphyte, 164

ANGIOSPERMAE — DICOTYLEDONEAE

PIPERACEAE

Peperomia tetraphylla (G. Forst.) Hook. & Am., forb, 72, 325

SALICACEAE

Salix capensis Thunb. var. capensis, tree, 295, 884

MYRICACEAE

Myrica

cordifolia L., shrub, 249

div ersifolia Adamson, shrub, 490

humilis Cham. & Schlechtd, shrub, 679

kraussiana Buchinger ex Meisn, shrub, 786

serrata Lam., shrub, 377, 837

ULMACEAE

Celtis africana Burm. f., tree, 110

Chaetacme aristata Planch., tree, 81, 293

MORACEAE

Ficus

burtt-davyi Hutch., tree, 962

ingens (Miq.) Miq. var. ingens, shrub or tree, 145, 705

sur Forssk., tree, 126, 1504

URTICACEAE

Laportea grossa (Wedd) Chew, forb, 1206, 1602

Didymodoxa caffra (Thunb.) Friis & Wilmot-Dear, forb, 1407

PROTEACEAE

Protea

cynaroides (L.) L., shrub, 319

foliosa Rourke, shrub, 98

lorifolia (Salisb. ex Knight) Fourc. , shrub or small tree, 258, 636

repens (L.) L., shrub, 1554

Leucospermum cuneiforme (Burm. f.) Rourke, shrub, 1, 614

Leucadendron salignum Berg., shrub, 41, 1278, 1325

VISCACEAE

Viscum

minimum Harv., epiphytic parasite, sight record

rotundifolium L. f., epiphytic parasite, 74

SANTALACEAE

Colpoon compressum Berg., small tree, 21, 202

Rhoiacarpos capensis (Harv.) DC., shrub, 1919

Thesium

disciflorum A. W. Hill, shrublet, 793, 1695

flexuosum A. DC., shrublet, 187, 609

fruticosum A. W. Hill, shrublet, 184

gnidiaceum A. DC., shrublet, 1716

junceum Bemh., shrublet, 832, 1296

pallidum A. DC., shrublet, 846

cf. squarrosum L. f., shrublet, 604, 1258

strictum Berg. , erect shrub, 97, 606, 1341

virgatum Lam., shrublet, 221, 353

POLYGONACEAE

Emex australis Steinh., forb, 1355*

226

Bothalia 18,2 (1988)

Rumex

crispusZ,., forb, 1915*

cf. nepalensis Spreng., forb, 2127*

cf. sagittatus Thunb., forb, 1932

Polygonum

aviculare L., forb, 1972*

lapathifolium L. subsp. maculatum (S.F. Gray) T.-Dyer& Trim.,

forb, 1937

salicifolium Willd., forb, 338, 936

Bilderdykia convolvulus (L.) Dumort, twining forb, 1916*

CHENOPODIACEAE

Chenopodium

carinatum R. Br., forb, 918, 1091*

mucronatum Thunb., forb, 1404

Exomis microphylla (Thunb.) Aell. var. axyrioides ( Fenzl ) Aell.,

shrub, 1261

Salsola kali L., forb, 1422*

AMARANTHACEAE

Am ar an thus

hibridus L. subsp. hibridus var. hibridus, forb, 1128*

thunbergii Moq., forb, 1405

Pupaha lappacea (L.) Juss., forb, 1370*

Achyranthes sicula (L.) All., forb, 1635*

Achyropsis

avicularis (E. Mey. ex Moq.) Hook, f., forb, 1082, 1615

leptostachya (E. Mey. ex Meisn.) Hook, f, forb, 981

Gomphrena celosioides Mart., forb, 1414*

AIZOACEAE

Limeum

africanum Burm. subsp. africanum, forb, 1373

telephioides E. Mey. ex Fenzl var. telephioides, forb, 905

Gisekia pharnaceoides L., forb, 1853

Pharnaceum dichotomum L. f, forb, 573, 1973

Galenia

pubescens (Eckl. & Zeyh.) Druce, shrublet, 1 744

sarcophylla Fenzl, shrublet, 1365

secunda (L. f.) Sond., shrublet, 1977

Aizoon

canariense L., forb, 1208

rigidum L. f. var. villosum Adamson, forb, 557

MESEMBRYANTHEMACEAE

Aptenia cordifolia (L. f.) Schwant. var. cordifolia, succulent, 993

1983

Bergeranthus

scapiger (Haw.) N.E. Br., succulent, 810

vespertinus (Berger) Schwant., succulent, 544

Carpobrotus dimidiatus (Haw.) L. Bol., succulent, 486, 612

Delosperma

brevisepalum L. Bol. var. brevisepalum, succulent, 572

cf. cooperi (Hook, f.) L. Bol., succulent, 2072 .

ecklonis (Salm-Dyck) Schwant., succulent, 1374

hirtum (N.E. Br.) Schwant., succulent, 819, 820

laxipetalum L. Bol, succulent, 543a

lehmannii (Eckl. dt Zeyh.) Schwant., succulent, 1967, 1976

cf. testaceum (Haw.) Schwant., succulent, 1354

sp., succulent, 1560

Lampranthus

productus (Haw.) N.E. Br., succulent, 1974

spectabilis (Harv.) N.E. Br. subsp. spectabilis, succulent, 697

1388

Mestoklema tuberosum (L.) N.E. Br. ex Glen, succulent, 814,

1733

Pleiospilos sp.?, succulent, 476

Ruschia

orientalis L. Bol., succulent, 502, 1331, 1542

stenophylla (L.) Bol.) L. Bol., succulent, 254, 1216, 1693

PORTULACACEAE

Anacampseros cf. telephiastrum DC., succulent, 266, 854, 1314

Portulacaria afra Jacq., large succulent, 526

CARYOPHYLLACEAE

Cerastium capense Sond., forb, 1925

Polycarpon tetraphyllum L. f., forb, 882, 1244, 1854, 1921*

Silene

burchellii Otth

var. angustifoha Sond., forb, 1308, 1 703

var. burchellii, forb, 1681

var. pilosellaefolia (Cham. & Schlechtd.) Sond., forb, 1183

gallicaZ., forb, 1671*

Dianthus zeyheri Sond. subsp. natalensis Hooper, forb, 1030

ILLECEBRACEAE

Polhchia campestris/4r'r., forb, 257, 921, 1035, 1438

RANUNCULACEAE

Knowltonia

cordata H. Rasm., forb, 1336, 1500, 2141

vesicatoria (L. f.) Sims subsp. vesicatoria, forb, 1692

Clematis brachiata Thunb., climber, 328

Ranunculus multifidus Forssk., forb, 330

MENISPERMACEAE

Cissampelos torulosa E. Mey. ex Harv., climber, 1899

LAURACEAE

Cassytha

ciliolata Nees, twining parasite, 91

filiformis L, twining parasite, 2165*

PAPAVERACEAE

Argemone subfusiformis G.B. Ownbey, forb, 911*

Papaver aculeatum Thunb., forb, 1958

FUMARIACEAE

Fumaria muralis Sond. ex Koch, twining forb, 1927

BRASSICACEAE

Heliophila

elongata (Thunb.) DC., forb, 1041, 1430

rigidiuscula Sond., forb, 360

trifurca Burch, ex DC., forb, 842

Lepidium africanum (Burm. f.) DC. subsp. africanum, forb, 1818

Coronopus didymus (L.) Sm., forb, 1866*

Sisymbrium

capense Thunb., forb, 1729

orientale L., forb, 1810*

Eruca sativa Mill., forb, 1595*

cf. Diplotaxis muralis (L.) DC., forb, 1776*

Raphanus raphanistrum L., forb, 1928*

Rorippa

fluviatilis (E. Mey. ex Sond.) Thell.

var. caledonica (Sond.) Marais, forb, 1815

var. fluviatilis, forb, 1 752

Capsella bursa-pastoris (L.) Medik., forb, 1124*

CAPPARACEAE

Capparis sepiaria L. var. citrifolia (Lam.) Toelken, tall shrub, 137,

140, 807

Boscia oleoides (Burch, ex DC.) Toelken, tree, 825

Cadaba aphylla (Thunb.) Wild, shrub, 1797

Maerua

cafra (DC.) Pax, tree, 752b

racemulosa (DC.) Gilg & Ben., tree, 66

DROSERACEAE

Drosera trinervia Spreng., forb, 1501, 1682

CRASSULACEAE

Cotyledon

campanulata Marloth, succulent, 930

cuneata Thunb., succulent, 1423 e

orbiculata L, succulent, 627, 718

velutina Hook, f., succulent, 761

woodii Schonl. & Bak. f, succulent, 79, 619, 1624b

Kalanchoe rotundifolia (Haw.) Haw., succulent, 133, 817, 1418

Crassula

arborescens (Mill) Willd. subsp. arborescens, succulent shrub,

618

capitella Thunb. subsp. thyrsiflora (Thunb.) Toelken, succulent,

708

cordata Thunb., succulent, 477, 858, 1787

ericoides Haw. subsp. ericoides, succulent, 467, 1306, 1432

expansa (Dryand.) Ait. subsp. expansa, succulent, 815, 1151

1638

lactea Soland., succulent, 52, 624, 1822

latibracteata Toelken, succulent, 640

Bothalia 18,2 (1988)

227

mesembryanthemoides (Haw.) Dietr.

subsp. hispida (Haw.) Toelken, 25, 1465

subsp. mesembryanthemoides, succulent, 812, 1565

muscosa L. cf. var. muscosa, succulent, 144

nemorosa (Eckl. & Zeyh.) Endl. ex Ml alp. , succulent, 76

nudicaulis L.

var. nudicaulis, succulent, 1 762

var. platyphylla (Harv.) Toelken, succulent, 49, 209, 796, 855

obovata Haw. var. obovata, succulent, 253, 1464

orbicularis L., succulent, 366, 473

ovata (Mill) Druce, succulent, 1641

pellucida L.

subsp. brachypetala (Drege ex Harv.)Toelken, succulent, 1578

subsp. marginalis (Dryand. in Ait.) Toelken, succulent, 1591,

1642

perfoliata L.

var. coccinea (Sweet) Rowley, succulent, 860a, 1230

var. minor (Haw.) Rowley, succulent, 1426, 1457

var. perfoliata, succulent, 1 783

perforata Thunb., succulent, 444, 540, 1626

pubescens Thunb.

subsp. radicans (Haw.) Toelken, succulent, 816

subsp. rattrayi (Schonl. & Bak. f.) Toelken, succulent, 1421,

156 7

rogersii Schonl., succulent, 1819, 2070

rupestris Thunb. subsp. rupestris, succulent, 37, 151, 260

tetragona L.

subsp. connivens (Schonl.) Toelken, succulent, 1804

subsp. lignescens Toelken, succulent, 1353, 1 772

subsp. robusta (Toelken) Toelken, succulent, 823

vaginata Eckl. & Zeyh. subsp. vaginata, succulent, 1530, 2104

sp. cf. C. capensis (L.) Baill., succulent forb, 1488

Adromischus

cristatus (Haw.) Lem. var. cristatus, succulent, 700, 1481

sphenophyllus C.A. Sm., succulent, 852, 1319

MONTINIACEAE

Montinia caryophyllacea Thunb., shrub, 181

PITTOSPORACEAE

Pittosporum viridiflorum Sims, tree, 150, 792

CUNONIACEAE

Cunonia capensis L., tree, 248

HAMAMELIDACEAE

Trichocladus ellipticus Eckl. & Zeyh. ex Walp., tree, 68, 287

ROSACEAE

Rubus

pinnatus Willd., scandent shrub, 750

rigidus Sm., scandent shrub, 1275

Alchemilla capensis Thunb., forb, 190, 2123

Agrimonia odorata Mill., shrub, 1242*

Cliffortia

burchellii Stapf, shrub, 185

ilicifolia L. var. ilicifolia, shrub, 409

linearifolia Eckl. & Zeyh., shrub, 199, 1305, 2168

paucistaminea Weim., shrub, 250, 421, 746

serpyllifolia Cham. & Schlechtd., shrub, 1502

strobilifera Murray, shrub, 32, 344, 1161

FABACEAE

Acacia

caffra (Thunb.) Willd., tree, 6 7

karroo Hayne, tree, 315

Schotia

afra (L.) Thunb. var. afra, tree, 542

latifolia Jacq., tree, 148, 1833

Calpurnea aurea (Ait.) Benth. subsp. sylvatica (Burch.) Brummitt,

small tree, 528, 1757

Podalyria burchellii DC., shrub, 7, 393, 598, 1433b

Rafnia elliptica Thunb., shrub, 615, 1066

Lotononis

cytisoides (E. Mey.) Benth., shrub, 494

pungens Eckl. & Zeyh., forb, 2196

Asp ala thus

biflora E. Mey. subsp. biflora, shrublet, 701

chortophila Eckl. & Zeyh .

subsp. chortophila, shrub, 101, 171, 988, 1118, 1436

subsp. kougaensis Dahlg., shrub, 468

frankenioides DC., shrub, 207, 428, 1290

lactea Thunb. subsp. adelphea (Eckl. & Zeyh.) Dahlg., shrub,

1564, 1831

laricifolia Berg. , shrub, 845

spinosa L. subsp. spinosa, shrub, 1303, 1440

teres Eckl. <6 Zeyh. subsp. teres, shrub, 431, 1329, 2149

Buchenroedera multiflora Eckl. & Zeyh., shrub, 1478, 1523

Melolobium canescens Benth., shrub, 927

Crotalaria capensis Jacq. , small tree, 1863

Argyrolobium

collinum Eckl. & Zeyh., forb, 645, 1348, 1441, 1535, 1856,

2042

crassifolium Eckl. & Zeyh., shrub, 464, 1522, 1838

molle Eckl. & Zeyh., forb, 1456, 1483, 2131

tomentosum (Andr.) Druce, shrub, 321, 1089, 1358b, 1607

tuberosum Eckl. & Zeyh., forb, 1447

sp. cf. A. pilosum Harv., forb, 1455, 1512, 2132

Medicago laciniata (L.) Mill., forb, 1364

Indigofera

acutisepala Conrath, shrub, 1063, 1894

denudata Thunb., shrub, 94, 234, 495

disticha Eckl. & Zeyh., shrub, 1420, 1619

fastigiata E. Mey., shrub, 1170

hedyantha Eckl. & Zeyh., shrub, 590, 1209, 2065, 2118

heterophylla Thunb., forb, 1349

sessilifolia DC., shrub, 1413

stenophylla Eckl. & Zeyh., shrub, 509, 834, 1326

stipularis Link, forb, 500, 600, 859, 126 7

stricta L. f., shrub, 1205, 1 700, 2164

sp. cf. I. brachystachya E. Mey., shrub, 251

Psoralea

asarina (Berg.) Salter, shrub, 1474, 1492

glabra E. Mey., shrub, 208, 1096, 1179, 149 7, 1581

pinnata L., shrub or small tree, 90, 785

Otholobium prodiens C.H. Stirton, shrub, 270, 774

Tephrosia

capensis (Jacq.) Pers.

var. acutiflora E. Mey., suffrutex, 747

var. angustifolia E. Mey., suffrutex, 152, 696, 1027

var. capensis, suffrutex, 2166

grandiflora (Ait.) Pers., shrub, 2154

macropoda (E. Mey.) Harv., shrub, 1051, 1169

semiglabra Sond., suffrutex, 2009

Sutherland^ frutescens (L.) R. Br., shrub, 2193

Lessertia cf. stenoloba E. Mey., forb, 1811

Zornia capensis Pers., forb, 111 7

Rhynchosia

caribaea (Jacq.) DC., climber, 327, 824, 1383

cooperi (Harv. ex Bak. f.) Bum Davy, climber, 374

totta (Thunb.) DC., forb, 1061, 1130, 1425, 1446, 1487

Eriosema salignum E. Mey., forb, 721, 861, 1039, 1207

Dipogon lignosus (L.) Verde., climber, 1191

GERANIACEAE

Geranium

caffrum Eckl. & Zeyh., forb, 2122

flanaganii Knuth, forb, 2155

incanum Bunn. f. var. multifidum (Sweet) Hilliard & Burtt,

forb, 386

Monsonia emarginata (L. f.) L’Herit., forb, 730, 116 7

Erodium cicutarium (L.) L’Herit. ex Ait., forb, 1809*

Pelargonium

alchemilloides (L.) L’Herit., forb, 758, 1064, 1358a

graveolens L ’Herit., forb ,51,329, 775

cf. grossularioides (L.) L’Herit., forb, 903

heracleifolium Lodd., forb, 1462

luridum (Andr.) Sweet, forb, 1310

multicaule Jacq. var. multicaule, forb, 390, 1529, 1699

odoratissimum (L.) L'Herit., forb, 1643

panduriforme Eckl. & Zeyh., forb, 915

peltatum (L.) L’Herit., creeper, 620, 1561

pulverulentum Colv. ex Sweet, forb, 1335, 1912

quercifolium (L. f.) L’Herit., forb, 913

reniforme Curt. , forb, 205, 396

ribifolium Jacq. , forb, 31, 613, 830

schizopetalum S weet, forb, 1139

sidifolium (Thunb.) Kunth, forb, 1285

zonale (L.) L’Herit., forb, 629

228

Bothalia 18,2 (1988)

OXALIDACEAE

Oxalis

bifi urea Lodd, geophyte, 1450, 1524, 2103, 2112

caprina L., geophyte, 1351

comiculata L., geophyte, 1743*

cf. setosa L., geophyte, 1454

smithiana Eckl. <6 Zeyh., geophyte, 1994

stellata EckL & Zeyh., geophyte, 1189

LINACEAE

Linum thunbergii Eckl. & Zeyh., forb, 397, 608, 1445, 1498

ZYGOPHYLLACEAE

Zygophyllum foetidum Schrad & WendL, shrub, 2195

RUTACEAE

Zanthoxylum capense (Thunb.) Harv., tree, 108, 159

Calodendrum capense (L. f.) Thunb., tree, 109

Agathosma

acutissima Duemmer, shrub, 416, 1552

capensis (L.) Duemmer, shrub, 1539

cf. peglerae Duemmer, shrub, 21 91

puberula (Steud.) Fourc., shrub, 567, 828, 1330

venusta (Eckl. & Zeyh.) Pillans, shrub, 269, 354

Coleonema pulchellum I. Williams, shrub, 564, 2001

Vepris lanceolata (Lam.) G. Don, tree, 136

Clausena anisata (Willd) Hook. f. ex Benth., tree, 1592

PTAEROXYLACEAE

Ptaeroxylon obliquum (Thunb.) Radik., tree, 142

MELIACEAE

Melia azedarach L„ tree, 1933*

POLYGALACEAE

Polygala

asbestina Burch., shrublet, 1385, 1774

fruticosa Berg. , shrub, 180, 414

hispida Burch., shrublet, 1204

illepida E. Mey., shrublet, 961, 1104, 1154, 1291, 1511

microlopha DC., shrublet, 1826

myrtifolia L., shrub, 157, 879, 1624a

uncinata E. Mey. ex Meisn., shrublet, 958, 1537

Muraltia

alopecuroides (L.) DC., shrublet, 280, 570, 850, 2163

ericifolia DC., shrub, 191, 426, 1484

EUPHORBIACEAE

Phyllanthus

burchellii Muell. Arg., forb, 950a

maderaspatensis L., forb, 950b

verrucosus Thunb., shrub, 113, 413

Croton rivularis Muell. Arg., tree, 53

Leidesia procumbens (L.) Prain, forb, 782

Acalypha

ecklonii Baill., forb, 907, 979, 1896, 1930

glabrata Thunb. var. glabrata, shrub, 54, 797

peduncularis E. Mey. ex Meisn., forb, 1 724

punctata Meisn., forb, 1125

Ctenomeria capensis (Thunb.) Harv. ex Sond., climber, 2010,

1358c

Dalechampia capensis Spreng. f., climber, 1834

Ricinus communis L., forb, 912*

Jatropha capensis (L. f.) Sond, shrub, 107

Clutia

affinis Sond., shrub, 38

alatemoides L., shrub, 172, 459, 492

ericoides Thunb., shrub, 474

heterophylla Thunb., shrub, 1040

pulchella L., shrub, 41 7

Suregada africana (Sond.) Kuntze, shrub or small tree, 1880

Euphorbia

heptagona L., succulent, 616

kraussiana Bemh. , forb, 104, 754

ledienii Berger, succulent, 527, 826

mauritanica L., succulent, 105

polygona Haw., succulent, 153

cf. pugniformis Boiss., succulent, 1288

silenifolia (Haw.) Sweet, geophyte, 48, 1 94

striata Thunb., forb, 659, 1580

tetragona Haw., succulent tree, 102

triangularis Desf, succulent tree, 631

Chamaesyce inaequilatera (Sond.) Sojak, forb, 883

ANACARDIACEAE

Harpephyllum caffrum Bemh. ex Krauss, tree, 33, 193

Schinus molle L., tree, 2194*

Loxostylis alata Spreng. f. ex Reichb., tree, 154

Rhus

cf. albomarginata Sond., shrub, rare, 866

chirindensis Bak. f forma legatii (Schonl.) R. & A. Fernandes,

tree, 63, 229

dentata Thunb. shrub, 36, 491, 1279

fastigiata Eckl. & Zeyh., shrub, 603, 771, 1298

incisa L. f. var. effusa (Presl) R. Fernandes, tree, 465

lancea L. f., tree, 928

longispina Eckl. & Zeyh., tree, 633, 939

lucida L. forma lucida, shrub, 463, 689

p aliens Eckl. & Zeyh., shrub, 120, 216, 243, 512

pyroides Burch, var. pyroides, tree, 61, 595

refracta Eckl. & Zeyh., tree, 158, 518, 558

rehmanniana Engl. var. uitenhagensis (Eckl. & Zeyh.) ined,

tree, 244

rigida Mill. var. rigida, shrub, 15, 740, 1453

rosmarinifolia Vahl, shrub, 169

AQUIFOLIACEAE

Hex mitis (L.) Radik., tree, 127

CELASTRACEAE

Maytenus

acuminata (L. f.) Loes. var. acuminata, tree, 14, 441

capitata (E. Mey. ex Sond.) Marais, shrub, 551, 1403

heterophylla (Eckl. & Zeyh.) N.K.B. Robson, shrub or tree,

223, 309, 442

linearis (L. f.) Marais, shrub, 2197

nemorosa (Eckl. & Zeyh.) Marais, tree, 56, 62, 316, 440, 1001

undata (Thunb.) Blakelock, tree, 523, 53 7

Putterlickia pyracantha (L.) Szyszyl, shrub, 111, 562, 635, 1402

Pterocelastrus tricuspidatus (Lam.) Sond, shrub or tree, 198, 513,

857, 1656

Cassine

aethiopica Thunb., tree, 246, 302, 534, 856, 1594, 1600

crocea (Thunb.) Kuntze, tree, 529

peragua L., shrub or tree, 85, 306, 446

reticulata (Eckl. & Zeyh.) Codd, shrub, 1115, 1555

tetragona (L. f.) Loes., scandent shrub, 300

Pleurostylia capensis (Turcz.) Oliv., tree, 77, 301, 517a, 1609

ICACINACEAE

Apodytes dimidiata E. Mey. ex Am. subsp. dimidiata, tree, 284

SAPINDACEAE

Allophylus decipiens (Sond) Radik., tree, 95, 457, 1 755

Atalaya capensis R.A. Dyer, tree, 949

Smellophyllum capense Radik., tree, 196

Pappea capensis Eckl. & Zeyh., tree, 132

Dodonaea angustifolia L. /, small tree, 565

Hippobromus pauciflorus (L. f.) Radik., tree, 122, 292

MELIANTHACEAE

Melianthus comosus Vahl, shrub, 879, 919

RHAMNACEAE

Scutia myrtina (Burm. f.) Kurz, woody climber, 149

Noltea africana (L.) Reichb. f., tree, 60

Phylica

axillaris Lam. var. microphylla (Eckl. & Zeyh.) Pillans, shrub,

4, 197,201,411,674, 1241

paniculata Willd., small tree, 273, 429

Helinus integrifolius (Lam.) Kuntze, climber, 1924

VITACEAE

Rhoicissus

digitata (L. f.) Gilg & Brandt, woody climber, 70, 117

tomentosa (Lam.) Wild & Drum., woody climber, 143

tridentata (L. f.) Wild & Drum., woody climber, 13, 333, 438,

589

sp., shrub, 1800

Cyphostemma cirrhosum (Thunb.) Desc. ex Wild & Drum., suc-

culent climber, 922

Both alia 18,2 (1988)

229

TILIACEAE

Grewia

occidentalis L., shiub or small tree, 103, 430, 451

robusta Burch., shrub or small tree, 545, 548

MALVACEAE

Abutilon sonneratianum (Cav.) Sweet, shrub, 556, 1073

Malvastrum coromandelianum (L.j Garcke, shrub, 1 731 *

Sida

dregei Burtt Davy, forb, 947

ternata L. f, forb, 559, 1078

Pavonia praemorsa (L. f.) Cav., shrub, 320, 1612

Hibiscus

aethiopicus L.

var. aethiopicus, forb, 1252

var. angustifolius (Eckl. <t Zeyh.) Exell, forb, 1021

pusillus Thunb., forb, 945, 1081

trionum L., forb, 1178

STERCULIACEAE

Melhania didyma Eckl. & Zeyh., forb, 960, 1080

Herman nia

althaeoides Link, shrublet, 605, 1136

flammula Harv., shrublet, 10

geniculata Eckl. & Zeyh., shrub, 1243

cf. hyssopifolia L., shrub, 895

cf. odorata Ait., shrub, 597

OCHNACEAE

Ochna

arborea Burch, ex DC. var. arborea, tree, 80

natalitia ( Meisn .) Walp., tree, 141, 308

serrulata (Hochst.) Walp., shrub, 379, 666

CLUSIACEAE

Hypericum

aethiopicum Thunb. subsp. aethiopicum, forb, 1302, 1528

lalandii Choisy, forb, 1282, 1494

VIOLACEAE

Hybanthus capensis (Thunb.) Engl., forb, 984, 1025

FLACOURTIACEAE

Scolopia

mundii (Eckl. & Zeyh.) Warb., tree, 135, 435

zeyheri (Nees) Harv., tree, 312, 755, 1648

Homalium

dentatum (Harv.) Warb., tree, new record for the Cape Province,

898, 1851

rufescens Benth., tree, 875, 1236, 1651, 1957

Trimeria trinervis Harv., tree, 34, 96, 1135

Dovyalis

rhamnoides (Burch, ex DC.) Harv., shrub, 298

zeyheri (Sond.) Harv., shrub, 155, 1114, 1647

ACHARIACEAE

Acharia tragodes Thunb., climber, 1756

CACTACEAE

Opuntia

aurantiaca Lindl., succulent forb, invasive, not collected*

ficus-indica (L.) Mill., succulent tree, invasive, not collected*

PENAEACEAE

Penaea cneorum Meerb. subsp. lanceolatum (L.) Dahlg., shrub,

252, 2004, 2170

OLINIACEAE

Olinia ventosa (L.) Cufod, tree, 227, 314, 870, 1620, 1630, 1632

THYMELAEACEAE

Gnidia

anthylloides (L. f.) Gilg., shrub, 43, 182, 1246

capitata L. f, shrub, 1009

coriacea Meisn., shrublet, 3

cuneata Meisn., shrub, 1412

sericea L. var. sericea, shrub, 1 714

thesioides Meisn. var. thesioides, shrub, 59, 571, 1380

Struthiola

argentea Lehm., shrublet, 12, 19, 173

cf. macowanii C.H. Wr., shrublet, 2041

parviflora Bartl. ex Meisn., shrublet, 655, 731

Passerina

falcifolia C.H. Wr., shrub, 2005, 2160

obtusifolia Thoday, shrub, 751

vulgaris Thoday, shrub, 83, 84

COMBRETACEAE

Combretum caffrum (Eckl.di Zeyh.) Kuntze, tree, 58, 524, 1423g

MYRTACEAE

Eugenia

capensis (Eckl. it Zeyh.) Harv. ex Sond., tree, 290, 764

zeyheri Harv., tree, 1134

ONAGRACEAE

Oenothera

indecora Cambess., forb, 1947*

parodiana Munz subsp. parodiana, forb, 1997*

HALORAGACEAE

Laurembergia repens Berg, subsp. brachypoda (Hiem) Oberm,

forb, 1686

Gunnera perpensa L., forb, 935

ARALIACEAE

Cussonia spicata Thunb. var. spicata, tree, 64

APIACEAE

Centella

affinis (Eckl. & Zeyh.) var. affinis, suffrutex, 241, 275, 1513

asiatica (L.) Urb., creeping forb, 245, 1299, 1576, 1922

glabra ta L. var natalensis/4 damson, suffrutex, 2007

virgata (L. f.) Drude var. virgata, suffrutex, 1327

sp., prostrate forb, 1262

Alepidea capensis (Berg.) R.A. Dyer var. capensis, forb, 607,

1058,1172

Torilis arvensis (Huds.) Link, forb, 787*

Conium sp., forb, 1 759, 1883, 1895

Lichtensteinia kolbeana H. Bol., forb, 1945

Heteromorpha trifoliata (Wendl.) Eckl. & Zeyh. , shrub, 24

Rhyticarpus difformis (L.) Benth. & Hook., shrub, 35, 1235

Ciclospermum leptophyllum (Pers.) Sprague, forb, 1926*

Pimpinella caffra (Eckl. & Zeyh.) Harv. , forb, Femiebrae plot 9. 28

Berula erecta (Hudson) Cov., forb, 346

Polemannia sp? cf. P. montana Schltr. & Wolff, shrub, rare, 458

Stenosemis caffra (Eckl. & Zeyh.) Sond., forb, 274, 1210

Peucedanum

capense (Thunb.) Sond.

var. capense, shrub, 42, 407

var. lanceolatum (E. Mey. ex Meisn.) Sond., shrub, 20, 188,

410

CORNACEAE

Curtisia dentata (Burm. f.) C.A. Sm„ tree, 28

ERICACEAE

Erica

adunca Benth., shrub, 654, 788

caffra L„ shrub or small tree, 683

cerinthoides L. var. cerinthoides, shrublet, 167

chamissonis Klotzsch ex Benth., shrub, 711, 835

copiosa Wendl var. copiosa, shrub, 189, 372, 1019, 1180, 1466,

2162

curviflora L. var. curviflora, shrub, 1459

deliciosa Wendl. f. ex Benth., shrub, 1237, 2136

demissa Klotzsch var. polyantha (Klotzsch ex Benth.) Dulfer,

shrub, 784, 1026

pectinifolia Salisb., shrub, 401, 1324

simulans Dulfer var. simulans, shrub, 5, 1111, 1342, 1544, 1553

unilateralis Klotzsch, shrub, 11, 720, 1540

sp., shrub, 445, 617

MYRSINACEAE

Myrsine africana L., shrub, 271, 484

Rapanea melanophloeos (L.) Mez, tree, 22

PRIMULACEAE

Samolus valerandi L., forb, 887

Anagallis arvensis L., forb, 794, 1071, 1914, 2063*

230

Bothalia 18,2 (1988)

PLUMBAGINACEAE

Plumbago auricula ta Lam., scan dent shrub, 532

SAPOTACEAE

Sideroxylon inerme L., tree, 282, 437

EBENACEAE

Euclea

crispa (Thunb.) Guerke var. crispa, tree, 16, 147, 1 70, 422

natalensis A. DC., shrub, 966

polyandra (L. f.) E. Mey. ex Hiem, shrub, 44, 195

schimperi (A. DC.) Dandy var. schimperi, tree, 89, 550, 1129,

1568

undulata Jacq. var. undulata, tree, 592, 6 78

Diospyros

lycioides Desf. subsp. lycioides, shrub or tree, 30, 505, 593,

628, 813

scabrida (Harv. ex Hiem) De Winter var. cordata (E. Mey. ex

DC.) De Winter, shrub, 304, 507, 625, 687

whyteana (Hiem) F. White, tree, 289

OLEACEAE

Chionanthus

foveolatus (E. Mey.) Steam subsp. foveolatus, small tree, 128,

447

peglerae (C.H. Wr.) Steam, small tree, 1194

Olea

capensis L. subsp. capensis, shrub, 1719

capensis L. subsp. macrocarpa (C.H. Wr.) Verdoom, tree, 92,

790

europaea L. subsp. africana (Mill.) P.S. Green, tree, 73, 423

woodiana KnobL, tree, 57

Jasminum

angulare Vahl, climber, 1411, 1758, 1917

cf. multipartitum Hochst., climber, 1419

SALVADORACEAE

Azima tetracantha Lam., scandent shrub, 93

LOGANIACEAE

Strychnos decussata (Pappe) Gilg, tree, 288

Nuxia floribunda Benth., tree, 283

Buddleja saligna Willd., shrub or small tree, 118

GENTIAN ACE AE

Sebaea hymenosepala Gilg, forb, 361, 11 74, 1448

Chironia

melampyrifolia Lam., forb, 712

tetragona L. f, forb, 848

Nymphoides thunbergiana (Griseb.) Kuntze, aquatic forb, 1589,

2030

APOCYNACEAE

Acocanthera oppositifolia (Lam.) Codd, shrub, 78

Carissa

bispinosa (L.) Desf. ex Brenan var. acuminata (E. Mey.) Codd,

shrub, 773

bispinosa (L.) Desf. ex Brenan var. bispinosa, shrub, 1004

haematocarpa (Eckl.) A. DC., shrub, 1423d

Gonioma kamassi E. Mey., tree, 47, 115, 1146

Pachypodium

bispinosum (L. f.) A. DC., succulent, 599, 2147

succulentum (L. f.) A. DC., succulent, 809

ASCLEPIADACEAE

Xysmalobium involucratum Decne., forb, 1704

Schizoglossum

atropurpureum E. Mey. subsp. tridentatum (Schltr.) Kupicha,

forb, 2071

cordifolium E. Mey., forb, 1094, 1685

Periglossum sp., forb, 1663

Pachycarpus dealbatus E. Mey., forb, 741, 868, 959

Asclepias

cancellata Burm. f, forb, 1309

crispa Berg., forb, 1175

expansa (E. Mey.) Schltr., forb, 1171

Cynanchum natalitium Schltr., climber, 648

Sarcostemma viminale (L.) R. Br., succulent climber, 652

Secamone

alpinii Schultes, climber, 139, 311, 1667

filiformis (L. f.) J.H. Ross, climber, 525, 1382

Sisyranthus sp., forb, 1684

Anisotoma pedunculata N.E. Br., forb, 1678

Stapelia sp., succulent, 727, 1534

Tylophora lycioides (E. Mey.) Decne., forb, 1367

Dregea floribunda E. Mey., climber, 530

CONVOLVULACEAE

Cuscuta campestris Yunck., twining parasite, 560, 1401 *

Falkia repens L. f, creeping forb, 1301

Convolvulus

bidentatus Bemh. apud Krauss, creeping forb, 1857

farinosus L., creeping forb, 886

Ipomoea

arachnosperma Welw., climber, new record for Cape Province,

1423a

simplex Thunb., forb, 2117

BORAGINACEAE

Ehretia rigida (Thunb.) Druce, shrub or small tree, 549

Cynoglossum hispidum Thunb., forb, 738, 1160

Lithospermum

cf. inomatum DC., forb, 901, 1116, 1307

scabrum Thunb., forb, 2140

VERBENACEAE

Verbena bonariensis L., forb, 1995*

Lantana rugosa Thunb., shrub, 952, 1688

Plexipus cuneifolius (L. f.) Rafin., shrublet, 2201

Priva meyeri Jaub. & Spach. var. meyeri, forb, 973

LAMIACEAE

Ajuga ophrydis Burch, ex Benth., forb, 1211, 1293

Teucrium

africanum Thunb., forb, 1812

trifidum Retz., forb, 942, 1131

Leonotis

leonurus (L.) R. Br., shrub, 27

ocymifolia (Burm. f.) Iwarsson var. ocymifolia, shrub, 419, 1507

Stachys

aethiopica L., forb, 1 771

graciliflora Presl, forb, 1369

scabrida Skan, forb, 382a

Salvia

aurita L. f. var. aurita, forb, 1226, 1280

repens Burch, ex Benth. var. repens, forb, 906, 1861

triangularis Thunb., forb, 504, 1835

Mentha longifolia (L.) Huds. subsp. capensis (Thunb.) Briq., forb,

318

Plectranthus

madagascariensis (Pers.) Benth.

var. aliciae Codd, forb, 1132

var. madagascariensis, forb, 1525

var. ramosior Benth., forb, 1604

spicatus E. Mey. ex Benth., forb, 1110, 1562, 1645

strigosus Benth., forb, 131

verticillatus (L. f.) Druce, forb, 385, 1234, 1375

SOLANACEAE

Nicandra physalodes Gaertn., forb, 2076*

Lycium

ferocissimum Miers, shrub, 938, 1423 f

oxycarpum Dun., shrub, 1805

Withania somnifera (L.) Dun, shrub, 925

Physalis peruviana L., forb, 1616*

Solanum

giganteum Jacq., shrub, 1599

gracile Dun, forb, 1 751

hermannii Dun, shrub, 76 7

nodiflorum Jacq. , forb, 1767

retroflexum Dun, shrub, 914, 1485, 1929

rigescens Jacq., shrub, 497, 757

tomentosum L., shrub, 920, 1556

Datura ferox L., forb, 1423c*

Nicotiana glauca R.A. Grab., small tree, 722, 1948*

SCROPHULARIACEAE

Verbascum virgatum Stokes, forb, 206 7*

Diascia capsularis Benth., forb, 638, 1444, 1526

Nemesia

cf. cynanchifolia Benth., forb, 1395

fruticans (Thunb.) Benth., forb, 893, 899, 1907

melissifolia Benth., forb, 1910

Bothalia 18,2 (1988)

231

Halleria lucida L., shrub or small tree, 75, 21 7

Freylinia undulata (L. f.) Benth., shrub, 392

Sutera

albiflora Verdoom, shrublet, 99, 381

argentea (L. f) Hiem, shrublet, 663

foliolosa (Benth.) Hiern, shrublet, 902

kraussiana (Benth.) Hiem, shrublet, 953

pinnatifida (Benth.) Kuntze, shrublet, 1415

roseoflava Hiern, forb, 355, 391, 634

Phyllopodium krebsianum Benth., forb, 1931, 1971

Dysanthes dubia (L.) Bernh., forb, 1045, 1508

Veronica anagallis-aquatica L., forb, 1 748

Alectra capensis Thunb., parasitic forb, 719

Graderia scabra (L. f.) Benth., forb, 1392, 2119, 2129

Buchnera dura Benth., forb, 1232, 1431, 1490, 1570

Cycnium tubulosum (L. f.) Engl, parasitic forb, 215, 1439

Striga bilabiata (Thunb.) Kuntze, parasitic forb, 1173

Harveya speciosa Bernh. ex Krauss, parasitic forb, 228b, 1333

Hyobanche sanguinea L., parasitic herb, 1855

SELAGINACEAE

Hebenstretia integrifolia L., shrublet, 1906

Selago

corymbosa L., shrub, 166

speciosa Rolfe, forb, 1346

Walafrida

geniculata (L. f.) Rolfe, shrublet, 1 749

saxatilis (E. Mey.) Rolfe, shrublet, 1372

witbergensis (E. Mey.) Rolfe, shrublet, 1966

BIGNONIACEAE

Tecomaria capensis (Thunb.) Spach, shrub, 531, 651, 1381

Rhigozum obovatum Burch., shrub, 1389

GESNERIACEAE

Streptocarpus

meyeri Burn, forb, 1334, 1449

rexii (Hook.) Lindl., forb, 566, 1075, 1888

ACANTHACEAE

Thunbergia

capensis Retz., prostrate shrublet, 822, 1168

dregeana Nees, prostrate shrublet, 941

Chaetacanthus sp. nov., shrublet, 224, 672, 1400

Ruellia cf. pilosa L. f, forb, 698

Crabbea nana Nees subsp. nana, forb, 368

Barleria obtusa Nees, shrublet, 538, 1551

Blepharis

capensis (L. f.) Pers.; shrublet, 818

integrifolia (L. f.) E. Mey. ex Schinz var. integrifolia, shrublet,

1739

cf. procumbens (L. f.) Pers., shrublet, 462

Peristrophe cemua Nees, shrublet, 533

Hypoestes forskaolii (Vahl) R. Br., shrublet, 130, 336, 1603

Siphonoglossa

leptantha (Lindau) ined.

subsp. late-ovata (C.B. Cl.) ined., shrublet, 1850

subsp. leptantha, shrublet, 1882

Isoglossa

bolusii C.B. Cl., shrublet, 1 794

grantii C.B. C7., shrublet, 1606

prolixa (Nees) Lindau, shrublet, 1893

Justicia capensis Thunb., shrub, 1113

PLANTAGINACEAE

Plantago

lanceolata L., forb, 1222*

virginica L., forb, 1 737*

RUB LACE AE

Kohautia amatymbica Eckl. & Zeyh., forb, 1666

Coddia rudis (E. Mey. ex Harv.) Verde., shrub or small tree, 160,

686

Gardenia thunbergia L. f, tree, 123

Hyperacanthus amoenus (Sims) Bridson, tree, 86

Canthium

inerme (L. f.) Kuntze, tree, 106, 310, 436

mundianum Cham. & Schlechtd., tree, 65, 1646

Psydrax obovata (Eckl. & Zeyh.) Bridson subsp. obovata, tree,

71, 299, 303, 685, 869

Pavetta

capensis (Houtt.) Brem. subsp. capensis, tree, 26, 100, 1150

inandensis Brem. , tree, 294, 305

lanceolata Ec kl., tree, 1601

Psychotria capensis (Eckl.) Vatke, tree, 87, 684

Galopina

aspera (Eckl. & Zeyh.) Walp., forb, 1133

circaeoides Thunb., forb, 1152, 1227

Anthospermum

galioides Reichb. f. subsp. galioides, shrublet, 1304

herbaceum L. f, forb, 256, 841

paniculatum Cruse, shrub, 1300, 1321, 2096

spathulatum Spreng.

subsp. spathulatum, shrub, 121

subsp. uitenhagense Puff, shrub, 192

Richardia humistrata (Cham. & Schlechtd.) Steud., forb, 1668,

2008*

Rubia petiolaris DC., forb, 1 725

DIPSACACEAE

Cephalaria humilis (Thunb.) Roem. & Schult., forb, 955, 1182,

2097

Scabiosa columbaria L., forb, 399, 956, 1345

CUCURBITACEAE

Zehneria scabra (L. f.) Sond., climbing forb, 1608

Kedrostis

cf. capensis (Sond.) A. Meeuse, climbing forb, 811

nana (Lam.) Cogn. var. zeyheri (Schrad.) A. Meeuse, climbing

forb, 1376

Coccinea palmata (Sond.) Cogn., climbing forb, 1 779

CAMPANULACEAE

Wahlenbergia

capillacea (L. f.) A. DC., forb, 222, 1495

cf. cuspidata V. Brehm., forb, 2152

cf. dentifera V. Brehm., forb, 1069, 1698

krebsii Cham, subsp. krebsii, forb, 2135

madagascariensis A. DC., forb, 1750, 1860

undulata (L. f.) A. DC., forb, 801, 804, 904, 1621, 1908

lightfootia

albens Spreng. ex A. DC., shrublet, 586

cinerea (L. f.) Sond., shrublet, 1536, 1538, 1558

divaricata Buek.

var. debilis (Sond.) Adamson., forb, Plots 27.41, 28.30

var. divaricata, forb, 985

LOBELIACEAE

Cyphia sylvatica Eckl. & Zeyh. var. sylvatica, twining forb, 203,

380, 1557

Lobelia

ancepsZ,. f, forb, 1070, 1649

erinus L., forb, 2038

flaccida (Presl) A. DC. subsp. flaccida, forb, 1031, 1166

thermalis Thunb., forb, 877, 1862

tomentosa L. f, forb, 356, 1077

Monopsis

scabra (Thunb.) Urb., forb, 50, 1072, 1532

unidentata (Dryand.) E. Wimm. subsp. intermedia ined. , forb,

1532, 1676

Grammatotheca bergiana (Cham.) Presl var. bergiana, forb, 2035

ASTERACEAE

Vernonia

capensis (Houtt.) Druce, forb, 1018, 1059

neocorymbosa Hilliard, shrub, 1463

Corymbium africanum L., forb, 2000

Mikania capensis DC., climber, 1605

Pteronia

cf. paniculata Thunb., shrub, 1023

teretifolia (Thunb.) Fourc., shrub, 165, 1281

Amellus strigosus (Thunb.) Less, subsp. pseudoscabridus Rommel,

forb, 1736

Aster

bakenanus Burtt Davy ex C.A. Sm., shrublet, 1012, 1350, 2120

squamatus (Spreng.) Hieron, forb, 1814*

Felicia

aethiopica (Burm. f.) H. Bol. & Wolley Dodd ex Adamson &

Salter subsp. ecklonis (Less.) Grau, shrub, 1527

fascicularis DC., shrub, 900, 1076

filifolia (Vent.) Burtt Davy, shrub, 643

Microglossa mespilifolia (Less.) B.L. Robinson, scandent shrub,

1147, 1417, 1563, 1674

Nidorella undulata (Thunb.) Sond. ex Harv., forb, 1231, 1588

Conyza

scabrida DC., forb, 779

ulmifolia (Burm. f.) Kuntze, forb, 204, 332, 373, 1219, 1583

232

Bothalia 18,2 (1988)

sp., forb, 1 753

Chrysocoma ciliata L., shiub, 226, 278, 726

Brachylaena

elliptica (Thunb.) DC., tree, 39, 286

glabra (L. f.) Druce, tree, 124

ilicifolia (Lam.) Phill. & Schweick., shrub or tree, 563

Tarchonanthus camphoratus L., tree, 23

Blumea cafra (DC.) O. Hoffm., forb, 1852

Denekia capensis Thunb., forb, 1687

Gnaphalium

coarctatum Willd., forb, 776, 1934*

pensylvanicum Willd., forb, 770*

vestitum Thunb., forb, 427

Troglophyton capillaceum (Thunb.) Hilliard & Burtt subsp. capil-

laceum, forb, 1371

Pseudognaphalium undulatum (L.) Hilliard & Burtt, forb, 916,

1 763b

Helipterum milleflorum (L.) Druce, shrub, 46, 702

Helichrysum

albanense Hilliard, forb, 8, 1 78

anomalum Less., shrublet, 1482

appendiculatum (L. f.) Less., forb, 1214, 1702

aureum (Houtt.) Men. var. monocephalum (DC. ) Hilliard, forb,

714

cephaloideum DC., forb, 259, 469, 498

cymosum (L.) D. Don subsp. cymosum, shrublet, 1016, 1295

felinum Less., shrublet, 210, 1015

gymnocomum DC., forb, 1256

herbaceum (Andr.) Sweet, forb, 6, 1477

lineare DC., shrublet, 908, 909

miconiifolium DC., forb, 1038, 1215, 1294

nudifolium (L.) Less., forb, 412, 543b, 1140

odoratissimum (L.) Sweet, shrublet, 276, 602, 840

oxyphyllum DC., forb, 968, 1015

pilosellum (L. f.) Less., forb, 2130

rosum (Berg.) Less.

var. arcuatum Hilliard, shrublet, 805, 1002, 1969

var. rosum, shrublet, 843, 910

rugulosum Less., shrublet, 466

rutilans (L.) D. Don, shrublet, Plots 32.12, 22.3

spiralepis Hilliard & Burtt, forb, 680, 84 7

zeyheri Less., shrublet, 1339

Disparago ericoides Gaertn., shrublet, 272, 371

Elytropappus rhinocerotis (L. f.) Less., shrub, 138, 1798

Metalasia

gnaphalodes (Thunb.) Druce, shrub, 29

muricata (L.) D. Don, shrub, 681, 1473

Relhania

genistifolia (L.) L 'Herit., shrub, 1991

pungens L ’Hdrit.

subsp. angustifolia (DC.) Bremer, shrub, 1024, 1137

subsp. pungens, shrub, 424

Athrixia heterophylla (Thunb.) Less., forb, 402, 2066, 2139

Pulicaria scabra (Thunb.) Druce, forb, 1610

Xanthium spinosum L., forb, 1 785*

Verbesina encelioides (Cav.) Gray var. exauriculata C.B. Robinson

& Greenm., forb, 1423b*

Bidens pilosa L., forb, 1358d, 1550*

Galinsoga ciliata (Raf.) Blake, forb, 1361, 2074b*

Schkuhria pinnata (Lam.) Cabr., forb, 1845*

Tagetes minuta L., forb, 1 732*

Eriocephalus africanus L., shrub, 568

Lasiospermum bipinnatum (Thunb.) Druce, forb, 1807

Oedera imbricata Lam., shrub, 21 74

Athanasia

dentata (L.) L., shrub, 239, 641, 106 7

pinnata L. f, shrub, 483

Inulanthera dregeana (DC.) Kallersjo, shrub, 425, 594, 1468,

1516

Hymenolepsis indivisa (Harv.) Kdllersjo, shrub, 853

Matricaria nigellifolia DC. var. tenuior DC., forb, 1574

Cotula

coronopifolia L., forb, 897

heterocarpa DC., forb, 261, 363, 756, 1074, 1913

Schistostephium crataegifolium (DC.) Fenzl ex Harv., shrub, 45,

479

Artemisia afra Jacq. ex Willd., shrub, 335

Pentzia

incana (Thunb.) Kuntze, shrub, 535, 1387

pilulifera (L. f.) Fourc., forb, 926

Cineraria lobata L’Hirit., shrub, 1406, 1627, 1637, 1654

Senecio

acaulis (L. f.) Sch. Bip., forb, 2106

articulatus (L.) Sch. Bip., forb, 878

brachypodus DC., shrub, 1829, 1900

chrysocoma Meerb., forb, 971

coronatus (Thunb.) Harv., forb, 1033

crassiusculus DC., forb, 1017

crenatus Thunb., forb, 1213, 1472

gramineus Harv., forb, 378

ilicifolius L., forb, 1 766

inaequidens DC., forb, 324, 382b, 781

inomatus DC., shrub, Plot 9

isatideus DC., forb, 1347

juniperinusE./ var.juniperinus, forb, disturbed places, 277, 957

leptophyllus DC., forb, 917

linifolius L., forb, 1569, 1839, 2124

lineatus (L. f.) DC., shrub, Ferniebrae plots C8 & C9

madagascariensis Poir., forb, 1763a, 1951

oliganthus DC., forb, 644, 1467

othonniflorus DC., forb, 954, 1011, 1217, 2109

oxydontus DC., forb, 384, 982, 1112, 1337

oxyriifolius DC., forb, 1103, 1181

pinifolius (L.) Lam., shrub, 653, 2121

cf. pinifolius (L.) Lam., shrub, 584, 1849

polyodon DC. var. subglaber (Kuntze) Hilliard & Burtt, forb,

176, 387, 1442

puberulus DC., forb ,225, 970, 976

pyramidatus DC., shrub, 2144

quinquelobus (Thunb.) DC., shrub, 1597

radicans (L. f.) Sch. Bip., succulent forb, 583

retrorsus DC., forb, 2095

ruwenzoriensis S. Moore, forb, 1769

scaposus DC., forb, 1799

stria tifolius DC., forb, 1832

vitalis N.E. Br., succulent shrub, 114, 622, 699

sp. cf. S. albanensis DC. var. doroniciflorus (DC.) Harv., forb,

Plot 12.6

Euryops

algoensis DC., shrub, 587

euryopoides (DC.) B. Nord., shrub, 2021

latifolius B. Nord, sparse shrub, subendemic, 487, 580

trilobus Harv. , shrub, 724

Othonna quinquedentata Thunb., shrub, 362

Osteospermum

calendulaceum L. f., forb, 1362, 1622

glabrum N.E. Br., forb, 2006

herbaceum L. f., forb, 1409

imbricatum L.

subsp. imbricatum, shrub, 1 74, 601, 829, 1343

subsp. nervatum (DC.) Norl var. nervatum, shrub, 682, 986

junceum Berg., shrub, 836, 1119

Cluysanthemoides monilifera (L.) Norlindh subsp. pisifera (L.)

Norlindh, shrub, 146

Ursinia anethoides (DC.) N.E. Br., shrub, 175, 388, 588, 2107,

2159

Arctotis microcephala (DC.) Beauv., forb, 389, 596, 610, 924

Haplocarpha

lyrata Harv., forb, 220, 279

scaposa Harv., forb, 1062, 2137

Gazania

krebsiana Less.

subsp. arctotoides (Less.) RoessL, forb, 211, 365

subsp. serrulata (DC.) RoessL, forb, 716

Cullumia decurrens Less., forb, 394

Berkeya

decurrens (Thunb.) Willd, forb, 1228

sphaerocephala (DC.) RoessL, forb, 1225

sp., forb, 931

Carduus tenuiflorus Curtis, forb, 759*

Centaurea melitensis L., forb, 800*

Oldenburgia grandis (Thunb.) Baillon, tree, 129

Gerbera

ambigua (Cass.) Sch. Bip., forb, 737, 1543, 1697

piloselloides (L.) Cass., forb, 1092, 1212, 1251, 1496

Tolpis capensis (L.) Sch. Bip., forb, 1297

Hypochoeris

brasiliensis (Less.) Griseb., forb, 1975*

radicata L., forb, 1935*

Sonchus

dregeanus DC., forb, 1270, 1672

gigas L. Boulos ex Humbert, forb, 1253

oleraceus L., forb, 1936*

Bothalia 18,2: 233-260 (1988)

A synopsis of the plant communities of Swartboschkloof, Jonkershoek,

Cape Province

d.j. McDonald* **

Keywords: classification, forest, fynbos, map, Mesic Mountain Fynbos, phytosociology, Swartboschkloof

ABSTRACT

Swartboschkloof forms part of the Jonkershoek catchment complex, Cape Province, South Africa. It was select-

ed for multidisciplinary studies of Mountain Fynbos vegetation in 1978.

In this study the Braun-Blanquet (B-B) phytosociological method was used to study the vegetation. Data were

collected at 201 sites throughout the study area; 101 of these sites were correlated with a survey of soils of part

of the same area. Sixteen fynbos communities are classified in three groups and five forest communities are classified

in two groups.

Syntaxonomic tables and a map of the plant communities are presented and an attempt has been made to ex-

plain the distribution of the communities in terms of environmental factors.

UITTREKSEL

Swartboschkloof vorm deel van die Jonkershoekopvanggebiedstelsel, Kaapprovinsie, Suid-Afrika. Dit is in

1978 gekies vir multidissiplinere studies van Bergfynbosplantegroei.

In hierdie studie is die Braun-Blanquet-fitososiologiese metode (B-B) gebruik vir die bestudering van die plan-

tegroei. Gegewens is by 201 plekke regdeur die studiegebied versamel; 101 van hierdie plekke is gekorreleer met

’n opname van die gronde van deel van dieselfde gebied. Sestien fynbosgemeenskappe word in drie groepe ge-

klassifiseer en vyf woudgemeenskappe word in twee groepe geklassifiseer.

Sintaksonomiese tabelle en ’n kaart van die plantgemeenskappe word gegee en daar word gepoog om die ver-

spreiding van die gemeenskappe in terme van omgewingsfaktore te verduidelik.

CONTENTS

Introduction 234

Study area 234

Location 234

Geology, geomorphology and topography .... 235

Soils 235

Climate 235

Methods 237

Vegetation 237

1 Mesic Mountain Fynbos Communities 240

1.1 Erica hispidula-Diospyros glabra Shrublands . 240

1.1.1 Diospyros glabra-Elegia capensis High Closed

Shrubland (A) 240

1.1.2 Diospyros glabra- Cliff ortia odorata High

Closed Shrubland (B) 241

1.1.3 Diospyros glabra-Rhus angustifolia Short to

Tall Closed Shrublands 242

1 .1 .3.1 Rhus angustifolia-Berzelia lanuginosa Short

Closed Shrubland (D) 245

1 . 1 .3 . 1 . 1 Berzelia lanuginosa-Merxmuellera cincta

Tall Closed Shrubland (C) 245

1.1.3. 2 Rhus angustifolia-Zantedeschia aethiopica

High Closed Shrubland (E) 246

1.1.3. 3 Rhus angustifolia- My rsine africana Short

to High Closed Shrubland 247

1.1. 3.3.1 Myrsine africana-Olea europaea subsp.

africana High Closed Shrubland (F) 247

1 .1 .3.3.2 Myrsine africana- Giff ortia dentata Short

Closed Shrubland (G) 248

* Based on an M.Sc. thesis, University of Cape Town.

** Botanical Research Unit, P.O. Box 471, Stellenbosch 7600.

MS. received: 1987.08.19.

1.1.3 A Rhus angustifolia- Ischyrolepis gaudichau-

diana High Closed Shrubland (I) 248

1 .1 .3.4.1 Ischyrolepis gaudichaudiana-Myrsine afri-

cana High Closed Shrubland (H) 248

1.2 Diospyros glabra-Protea repens Transitional

Shrublands 249

1.2.1 Protea repens-Rhus angustifolia High Closed

Shrubland (L) 249

1.2.1 .1 Rhus angustifolia-Ischyrolepis sieberi Tall

Closed Shrubland (K) 250

1.2. 1.2 Protea repens-Maytenus oleoides High

Closed Shrubland (J) 250

1.2.2 Protea repens-Nebelia paleacea High Closed

Shrubland 251

1 .2.2.1 Nebelia paleacea-Erica sphaeroidea High

Closed Shrubland (M) 251

1.2. 2. 2 Nebelia paleacea- Restio perplexus Low

Closed Shrubland (N) 252

1 .3 Erica hispidula-Ischyrolepis sieberi Shrublands 253

1.3.1 Ischyrolepis sieberi-Nebelia paleacea Short

Closed Shrubland (P) 253

1.3.2 Ischyrolepis sieberi-Tetraria involucrata

Short Closed Shrubland (R) 253

2 Riparian and Forest Communities 254

2.1 Hartogiella schinoides-Diospyros glabra Ripa-

rian and Forest Communities 254

2.1.1 Halleria elliptica-Brabejum stellatifolium

Short Forest (T) 255

2.1.2 Halleria elliptica-Giff ortia cuneata High

Closed Shrubland (S) 255

2.2 Diospyros glabra-Rapanea melanophloeos Tall

Forest (W) 256

2.2.1 Rapanea melanophloeos- Cunonia capensis

High Forest (U) 256

234

Bothalia 18,2 (1988)

2.2.2 Rapanea melanophloeos-Heeria argentea

Short Forest (V) 257

Discussion and conclusions 257

Acknowledgements 259

References 259

Appendix 259

INTRODUCTION

This paper is a summary of information and data

gathered for a thesis (McDonald 1983) and previously

partly published in a report of the South African National

Scientific Programmes (McDonald 1985). A checklist of

the plants recorded from Swartboschkloof is published

separately (McDonald & Morley 1988).

Swartboschkloof is situated in the Jonkershoek Forest

Reserve, about 15 km from Stellenbosch in the Cape

Province. It is 373 ha in extent and forms part of the

greater Jonkershoek Catchment in the Hottentots-

Holland Mountains at 34° 00' south latitude and 18° 57'

east longitude. Most of the area has an equatorial (north-

facing) aspect.

Three studies of the vegetation of Swartboschkloof

(Van der Merwe 1966; Werger, Kruger & Taylor 1972;

Van Wilgen 1981) were carried out prior to this survey.

Those of Van der Merwe and Van Wilgen were mainly

concerned with vegetation response to fire and that of

Werger et al. was a phytosociological study. Fry (1987)

surveyed the soils of Swartboschkloof.

The objectives of this study were: a, to identify, de-

scribe and classify the Cape fynbos and forest communi-

ties occurring in Swartboschkloof; b, to map the plant

communities of Swartboschkloof; c, to relate the plant

communities to the soils of Swartboschkloof; d, to re-

late the plant communities to the abiotic environment.

STUDY AREA

Location

The boundaries of the area selected for this survey

(Figure 1) do not coincide exactly with the boundaries

of the Swartboschkloof-Sosyskloof Nature Reserve (pro-

claimed in 1936) but are as follows: in the north-east

FIGURE 1. — Aerial photograph of Swartboschkloof showing the main features and boundaries of the study area.

Bothalia 18,2 (1988)

235

the boundary is a ridge of sandstone cliffs; from the

south-east corner to the south-west corner the firebreak

forms the southerly boundary; on the west side the

boundary is the watershed between the Swartboschkloof

and Sosyskloof catchments; in the north, Sosyskloof

stream forms the boundary; a short firebreak bounds a

small section at the base of Swartboschkloof on the

north to north-east side.

At present Swartboschkloof is managed as part of the

whole Jonkershoek catchment complex.

Geology, geomorphology and topography

Rust (1967) classified the sandstone sediments of

Jonkershoek in the Graafwater Formation of the Table

Mountain Group (T.M.G.) and described the Stellenbosch

Mountain, out of which Swartboschkloof is eroded, as

‘a succession of mainly white or cream orthosandstone

with interlaminated purple siltstone beds, which show

some worm remains’. The Table Mountain Group lies un-

conformably on porphyritic Cape granite at Jonkershoek

(Taljaard 1949). Van der Merwe (1966) noted that the

granite is encountered up to about 530 m elevation above

which T.M.G. sandstone occurs. Fry (1987), however,

found the highest point of the granite-sandstone contact

to be nearer 700 m elevation.

Swartboschkloof is a fan-shaped valley ranging from

285 m to 1 200 m in altitude. The head of the valley

forms the widest part of the ‘fan’ and it narrows with a

decrease in altitude. The slopes range from less than 5°

to 45° with the steep slopes averaging 30° . About two

per cent of the area consists of inaccessible, almost verti-

cal cliffs (Figure 2).

Apart from the steep cliffs and colluvial slopes there

are also loose boulder screes. These screes consist of sand-

stone boulders and are on the higher slopes immediately

below the steep cliffs. In parts the screes have stabilized

and support forest vegetation.

The streams in Swartboschkloof follow the fault lines.

The main Swartboschkloof stream, which flows from

south-west to north-east, is perennial and well wooded.

It is fed by three other seasonal streams. The other main

stream, on the east side of the valley, is perennial but is

not fed by other streams.

Soils

The soils of Swartboschkloof are complex owing to

colluvial mixing of sandstone and granite. The presence

of granite gives rise to soils of the Hutton, Magwa and

Nomanci Forms. Seeps and other areas of impeded drain-

age are also found, resulting in development of soils of

the Westleigh and Fernwood Forms.

Fry (1987) classified the soils of Swartboschkloof

following the binomial classification for soils in South

Africa (MacVicar et al. 1977). He identified 12 forms

with 14 series.

Climate

Wicht, Meyburgh & Boustead (1969) described the

climate of Jonkershoek as being ‘Etesian of the Mediter-

ranean type’. According to Koppen’s (1931) system it

would have an average temperature of below 22° C for

the warmest month. It conforms to Climate Type IV in

Walter & Leith’s classification (Werger et al. 1972).

Swartboschkloof, having a general equatorial aspect,

experiences high insolation over the greater part but parti-

cularly on the higher slopes in the summer. The few slopes

with a southerly aspect experience less direct sun and are

thus cooler and moister.

Wicht et al. (1969) discussed at length the occurrence

of rain and its measurement, temperature changes and

incoming radiation. The mean annual rainfall over 20

years is about 1 600 mm at the base of the Swartbosch-

kloof Valley (Swartboschkloof weather station), 50%

of which falls from May to August and only 12% from

December to March — the hottest driest period.

FIGURE 2. — Swartboschkloof in

1973. Photograph by B . Bre-

denkamp, used with permis-

sion Jonkershoek Forestry

Research Centre.

236

Bothalia 18,2 (1988)

FIGURE 3. — Sketch-map of Swartboschkloof showing positions of releves and soil pits.

Bothalia 18,2 (1988)

237

METHODS

The Braun-Blanquet (B-B) approach was adopted in

this study. For the naming of the communities a species-

binomial system using two differential species in the syn-

taxon is used. Generally the first species was selected on

the basis of its dominance. No suffixes such as -etosum,

-etum and so on, which indicate syntaxonomic hierarchi-

cal ranking, have been used. Much more extensive sam-

pling of Mountain Fynbos is necessary before broad hier-

archical ranking and hence broad syntaxonomic nomen-

clature can be meaningfully applied to the communities

identified in this study (Barkman et al. 1986; S.A. Syn-

taxonomic Nomenclature Committee; Werger 1974). In

addition to the species-binomial, a structural name fol-

lowing Edwards (1983) is given for each community.

Taking the work of Werger (1972) and Campbell &

Moll (1981) into consideration, a pragmatic approach

was adopted where rectangular quadrats of 5 x 10 m

(50 m2) were used in the fynbos vegetation and 10 x

20 m (200 m2 ) quadrats in the riparian and forest com-

munities of Swartboschkloof. Quadrats of 50 m2 were

used to sample the Halleria elliptica-Brabejum Stella ti-

folium Short Forest as this size was adequate for this

community. In the case of the boulder scree communi-

ties, quadrats of 200 m2 were used; their shape was

dependent on subjective assessment of each stand.

The quadrats were divided, where possible, to give

10 equal-sized subplots, for systematized data record-

ing. The subplots were not nested. The long axis of each

quadrat of the fynbos samples was oriented parallel with

the contour whereas the long axes of the forest samples

had a downslope orientation.

There were two phases of fieldwork in this survey.

During the first phase, 101 of the 114 soil pits sampled

by Fry (1987) in his study of Swartboschkloof soils

were relocated. Vegetation sample plots were placed as

close as possible to the soil pits, to permit correlation

between the soil series identified (Fry 1987) and the

vegetation samples (Figure 3). This was done bearing in

mind the requirement for sampling ‘homogeneous’ stands

of vegetation (Werger 1974). It was, however, not always

possible to meet this requirement. The reason for this

was that the soil pits themselves were often on bounda-

ries in the soil catena, a feature reflected by the vegeta-

tion in some cases.

For the second phase of sampling it was necessary to

first examine and interpret 1:5 000 black and white and

1:20000 colour aerial photographs. This was done since

Fry (1987) did not cover the whole area, as defined for

this study, for his soil survey. A further 100 releves were

sampled during the second phase of fieldwork in the

vegetation units delimited on the aerial photos.

Floristic data were gathered by a, recording all per-

manently recognizable species together with their

cover-abundance (B-B) values; and b, collecting those

species not immediately recognizable. When ephemeral

species such as annuals and geophytes were encountered

they were recorded in parentheses.

Structural data were collected for each releve by re-

cording total cover (% ) for each stratum and the domi-

nant species in each stratum.

The computer program, TABSORT, developed by the

Directorate of Forestry at Jonkershoek and modified by

the Botanical Research Institute (Boucher 1977) was

used for the sorting of releve data in this study.

Tables 1, 2 and 3 are partial tables of the final fynbos

vegetation table. Summaries showing the relationships

between the communities represented in the respective

partial tables and the other plant communities are ap-

pended to each partial table. Map symbols are used for

simplicity to compare the summaries with the partial

tables. Table 4 summarizes the occurrence of the more

widely occurring species in the fynbos. Table 5 is the

final phytosociological table (with summary) of the

forest and riparian communities of Swartboschkloof

(McDonald 1983).

Geophytes and annuals which are not permanently

recognizable are not included in the tables. Species

with single occurrences or low cover-abundance are

excluded from the tables.

Original data for the Werger et al. (1972) survey were

made available to me by Mr H.C. Taylor. By keeping the

same releve order as in the final table of the Werger et al.

(1972) study and using the order of species from the

final tables of this study, it was possible to prepare B-B

tables from the Werger et al. (1 972) data which are direct-

ly comparable with those of this study. These tables are

not included here but the interpolation of the Werger

et al. (1972) releves is given in the plant community

descriptions presented by McDonald (1985).

Values in the phytosociological tables (Tables 1, 2, 3

& 5) follow the B-B cover-abundance scale (Mueller-

Dombois & Ellenberg 1974), with ‘O’ representing the

occurrence of species within 1 m around the outside of

any given releve. The values in the summaries and Table

4 represent the percentage occurrence of a species in the

group of releves representing each community as follows:

5 =80-100% ; 4 = 60-79% ; 3 =40-59% ; 2 =20-39% ;

1 =5-19%; +=1-5%.

The vegetation map (Figure 4) was prepared using an

enlargement (to 1:5 000) of a black and white aerial

photograph (Figure 1) as the base map. The photo was

annotated using stereoscopic viewing of 1 :20000 colour

aerial photographs for detailed information. The positions

of releves were plotted and a base map with tentative

community boundaries was drawn. The community

boundaries were then checked in the field, after which

the final map was prepared. Communities are indicated

by the letters A-W on the map and in the headings in

the text.

VEGETATION

The vegetation of Swartboschkloof has been divided

into two main types: 1, Mesic Mountain Fynbos ( sensu

Moll et al. 1984) and 2, Riparian and Forest Communi-

ties. This division is based on physiographic features as

well as physiognomic, structural and floristic character-

istics. The Mesic Mountain Fynbos (MMF) is the domi-

nant type and is found in drier situations than the forests

and riparian vegetation.

238

Bothalia 18,2 (1988)

FIGURE 4. — Vegetation map of Swartboschkloof. Symbols of plant communities axe explained in the headings in the text.

239

Bothalia 18,2 (1988)

TABLE 1.- Partial Braun-Blanquet (B-B) table (with summary table) of the flesic mountain Fynbos:

Erica hispidula - Diospyros glabra Shrublands

1.1.1 Differential species of the Diospyros glabra - Elegia capensis Shrubland

1 .1 .2 Differential species of the Diospyros glabra - Cliffortia odorata Shrubland

240

Table 1. continued...

Bothalia 18,2 (1988)

Community ref.no.

The MMF fits broadly into Taylor’s (1978) zonation

patterns. On the lower slopes with mixed granite-sand-

stone soils mainly proteoid vegetation occurs. On the

upper, drier slopes with shallow sandy soils, plants in

the families Restionaceae and Cyperaceae dominate.

Following Campbell’s (1985) classification, the proteoid

vegetation fits into the Jonkershoek Mesic Proteoid Fyn-

bos and the restioid-cyperoid vegetation approximates

the Kouga Mesic Restioid Fynbos. In addition localized

perennial seeps occur where typical hygrophilous com-

munities are found. These would be classified as Keurbos

Wet Ericaceous Fynbos in Campbell’s (1985) system.

The forests are restricted to streambanks of perennial

streams and rock screes and are somewhat shorter than

those of the southern Cape, probably because of more

extreme climatic conditions.

The Halleria elliptica-Brabejum stellatifolium Short

Forest grades into the high forest vegetation; it is most

often found along drainage lines which have seasonal

running water where high forest vegetation does not

develop. The plant communities described are based on

floristic data presented in Tables 1, 2, 3, 4 and 5.

The Mesic Mountain Fynbos communities are describ-

ed first, followed by descriptions of the forest and ripa-

rian communities. The order in which the communities

are described follows the order in which they are pre-

sented in Tables 1, 2, 3 and 4.

Three MMF groups, one forest and one riparian group

occur in Swartboschkloof. They are systematically sub-

divided on the basis of floristic relationships (Tables 1,

2, 3, 4 and 5). Map symbols are given in parentheses

with each community name to permit identification of

communities on the vegetation map (Figure 4).

1 Mesic Mountain Fynbos Communities

Three major Mesic Mountain Fynbos Communities

occur in Swartboschkloof: Erica hispidula-Diospyros

glabra Shrublands; Diospyros glabra-Protea repens

Transitional Shrublands and Erica hispidula-Ischyro-

lepis sieberi Shrublands.

1 . 1 Erica hispidula-Diospyros glabra Shrublands

The shrublands grouped together here (Table 1) in-

clude specialized communities and those communities

which favour heavier soils of granite or colluvial origin.

The effect of perennial free water appears to override

the influence of substrate on the development of the

Diospyros glabra-Elegia capensis Shrubland and the

Diospyros glabra- Cliff or tia odorata Shrubland. In this

sense, they are regarded as specialized communities to-

gether with the Rhus angustifolia-Berzelia lanuginosa

and Berzelia lanuginosa-Merxmuellera cincta Shrublands.

The latter two communities, however, are apparently

more substrate-dependent: they are found in a bottom-

land situation on deep, predominantly sandstone-derived

soils. These communities are all referred to as ‘hygro-

philous communities’.

The other shrublands differ from those outlined

above in their preference for drier situations on soils

where granite is the major parent material.

1.1.1 Diospyros glabra-Elegia capensis High Closed

Shrubland (A)

Campbell et al. (1981) structural equivalent: Tall Mid-dense

Shrubland

Bothalia 18,2 (1988) 241

TABLE 2.- Partial Braun-Blanquet ( B-B ) table (with summary) of the Mesic Mountain Fynbos: the Diospyros glabra - Protea repens Transitional Shrublands

Community ref. no. . 1.

(read vertically) . 1.

3.

. A

1 .

1 .

1

1 .

1

1 .

2. . Surrmary table

2.

2

Map symbol

. ABCDEF *GH . I JKLMNPR . ST*UVliJ .

Relev/e number

(read vertically)

. 1. 11111111

.666777777808899999 . 4444458888003555561 1.11111 2222233 .

.01 81 237891 23825785.1 5678245670383567534.1 578901 23736.

111. 11111111111. 1111111111111111.

44445566333 . 255502222334557 . 22223445777777889 .

03490557567 . 01 344235601 9295 . 64899274036789091 .

Species common to Communities 1.1. 3. 4 , 1.1. 3. 4.1 , 1.2.1 , 1.2. 1.1

212+11 +11+2021122

Ischyrolepis gaudichaudiana.2+1 1 21 ++1 +1 1 1 1

Oxalis versicolor .RR +++ ++R R RR

Pelargonium myrrhifolium

Otholobium spicatum . +1 + 01+11

Euphorbia genistoides

Helichrysum zeyheri

Phylica spicata

Pentaschistis aspera

Tetraria flexuosa

Agathosma juniperifolia

Metalasia cephalotes

Mohria caffrorum

Rhynchosia totta

Helichrysum teretifolium

Muraltia heisteria

Helichrysum auriculatum

Osteospermum tomentosum

Otholobium rotundifolium

Agathosma capensis

Crassula pruinosa

5 5.55+

4.41

3.2122

3.3+

3.2+

1 .12+21

2.2

.+11

. 12

Species common to Communities 1.1. 3.1, 1.1. 3.1

1, 1.1. 3. 2, 1.1. 3. 3.1, 1.1. 3. 3. 2, 1.3.4, 1.3. 4.1, 1.2.1, 2.1.1, 1.2. 1.2 4 1.2.2

Rhus angustifolia

Pteridium aquilinum

Anthospermum aethiopicum

Erica sphaeroidea

Maytenus oleoides

Rhus rosmarinifolia

Protasparagus rubicundus

Ficinia filiformis

Halleria elliptica

Chironia baccifera

Phylica pubescens

Agapanthus africanus

Helichrysum cymosum

Helichrysum odoratissimum

Aristida junciformis

Senecio umbel latus

Myrsiphyllum asparagoides

Chasmanthe aethiopica

Helichrysum crispum

0+++1+ 11 1 1+ +

+ 13534432 2132

0 111+11 ++1 +11 +

++ 111 +1+1 + ++

++0+ 00+ + +22

1 2++01 0++++1 +

+0+ + +1

+ 1 +0

2+141 3

+011 1 1 11101 +

+1 +

13 2

.1 R +++ R+

1 1+1

54245545.43331 .55 2

2245553.42311 .543 3

3445555.422+2 1+.5

413555.42313 + .4

25 35.4312 +♦+. 433145

12 23.43+21 ++ .3

135 4.21231 + .43

2 22.32221 + .2

3 15 3.2+ + .443

12 3.1 + + .3

2 521.212+1

551. +2 21 h

2 2 2+.111

1 3+. + + +

4 .++1

2 2 .1

15 .+ +

1 .+

+ .3 3

Species common to Communities 1 .1.3.1, 1.1. 3. 2, 1.1. 3. 3.1, 1.1. 3. 3. 2, 1.1. 3. 4, 1

1.3. 4.1, 1.2.1, 1.2. 1.1, 1.2. 1.2, 1.2. 2.1 4 1

Protea nitida

Protea repens

Brunia nodiflora

Montinia caryophyllacea

Themeda triandra

Peucedanum gummiferum

1 11 11 11.2 210 +221

11022+ 101 1 .042 +5 25 152

0 + 321. 220101 2

1 +12 1+111 ++ .0+ +1 ++ +++1+ +01 h

11 00 121.0 1 +

01444133323. 421 00

2 1110 0 . 21 1

+ ++ + 21. + ++

+ 1 01 . ++

244+

12+0

32 4535.222422 1.3

11 2.33 522+ .

11 +.22 32311.

512 5.443322 +.3

25 3.212211 .

115 2.1++ +++ .2

Differential species: Blechnum punctulatum, Crassula coccinea,

Crassula pellucida, Cullumia setosa, Elegia capensis, Hippia

pilosa, Lampranthus deltoides, Nemesia acuminata

Dominant species: Arctotis semipapposa, Blechnum punctula-

tum, Cullumia setosa, Elegia capensis, Psoralea pinnata

This Community is found on localized wet sites on

slopes at high altitudes (800—930 m) with a south-easterly

aspect, in the south-west quarter of the study area. The

soil is usually waterlogged. The Community is restricted

to this part of the area although many species which

occur in it are found in seep conditions at lower altitudes

associated with other species which do not occur here.

The estimated total cover of this Community is 98%.

Three strata can be distinguished. The tallest stratum

consists of shrubs and restios, 2—3 m high, with an

average projected canopy cover of 56% . Elegia capensis

and Psoralea pinnata are the dominant shrubs in this

stratum. Giffortia dentata, C. polygonifolia and Cullumia

setosa dominate in the second stratum which is 0,5— 2,0

m high and has an average projected canopy cover of 46% .

The third stratum (<0,5 m) consists of herbs and dwarf

shrubs and is dominated by Arctotis semipapposa and

Blechnum punctulatum. Projected canopy cover of the

low stratum is estimated at 10% on average.

1.1.2 Diospyros glabra-Giffortia odorata High Closed

Shrubland (B)

Campbell et al. (1981) structural equivalent: Low Closed Shrub-

land

Differential species: Centella eriantha, Giffortia odorata. Leu-

cadendron salicifolium

Dominant species: Giffortia odorata, Leucadendron salicifolium

242

Bothalia 18,2 (1988)

TABLE 3. Partial Braun-Blanquet (B-B) table (with summary) of the Mesic Mountain Fynbos:

Erica hispidula - Ischyrolepis sieberi Shrublands

Cliffortia exilifolia

The Diospyros glabra- Cliffortia odorata Community

is characterized on the basis of one releve (193), sampled

at the top of a stabilized scree slope below cliffs at an

altitude of 655 m. It occurs as a localized stand too small

to sample more extensively but for the sake of complete-

ness it is described. The distinction between it and the

other hygrophilous communities is shown in Table 1.

The Diospyros glabra- Cliffortia odorata Shrubland has

some species which are common to it and to the other

hygrophilous communities: Gnidia oppositifolia, Os-

mi topsis asteriscoides and Psoralea pinnata.

A dense growth of prostrate Cliffortia odorata domi-

nates the lower stratum (< 1 m). Emergent through this

layer are tall to high shrubs such as Leucadertdron sali-

ci folium, Osmitopsis asteriscoides and Psoralea pinnata,

with a projected canopy cover of 24% .

1.1.3 Diospyros glabra -Rhus angustifolia Short to Tall

Closed Shrublands

The common environmental factor shared by the sub-

categories in the Diospyros glabra-Rhus angustifolia

Shrublands is apparently the presence of granite or gra-

nite-derived material in the soil. Fry (1987) found granite

in 82% of the soil profiles related to these shrublands. It

is possible that the influence of granite is masked by an

overburden of sandstone-derived soil in the remaining

18% of the profiles.

The highest point of the granite-sandstone contact is

at about 700 m. Ten per cent of the releves representing

these shrublands are at higher altitudes (680-700 m)

near the granite-sandstone contact. The other 90% are

mostly situated at lower altitudes (330-550 m) where

Bothalia 18,2 (1988)

243

TABLE A.- Summary table of species with wide occurrence in the Mesic Mountain Fynbos of Swartboschkloof

244

Bothalia 18,2 (1988)

TABLE 5.- Braun-Blanquet (B-B) table of the forest communities of Suartboschkloof ,

with a summary table

Community ref. no. .2. . 2. . . 2. . 2. . 2. . Summary table

(read vertically) .1. . 1 . . . 2. . 2. . 2

.2 . 1 . . 1 . 2 .

flap symbol

s t * u v w .abcdef»hijkl™pr.st*uvw.

Releve number . 1.1111.11.1111222.1111.1111111.

(read vertically) . 3996.0001.16.6699000.1199.1966698.

. 3565346 . 7890 . 47 . 344901 2 . 5667 . 3301 283 .

2.1.2 Differential species of the Halleria elliptica - Cliffortia cuneata Shrubland

2.2.1 Differential species of the Rapanea melanophloeos - Cunonia capensis Forest

Species common to Communities 2.1.1, 2.1.2 S 2.2.2

Brabejum stellatifolium

Oxalis lanata

Rubus rigidus

Blechnum australe

.211 4.5555. . 3333.

. ++++.

. +111 . 4 .R2. 2212111. +

2 + .35 3 .

43353343322322.3 1 .

15 .21.

215 .33552 .

2.2.2 Differential species of the Rapanea melanophloeos - Heeria argentea Forest

Heeria argentea . ... .331+. . . 5 .

Bothalia 18,2 (1988)

245

TftBLE 5. continued...

Community ref. no. .2. . 2. . . 2. . 2. . 2. . Summary table

(read vertically) .1. . 1. . . 2. . 2. . 2.

.2 . 1 . . 1 . 2 .

Map symbol S T * U U UJ .ABCDEF*HI JKLmNPR.ST*UUlil.

Releve number . 1.1111.11.1111222.1111.1111111.

(read vertically) . 3996.0001.16.6699000.1199.1966698.

. 3565346.7890 . 47 . 344901 2 . 5667 . 3301 283 .

Species common to Communities 2.2, 2.2.1 4 2.2.2

Rapanea melanophloeos

Olinia ventosa

1323211

1+1 +++

1 +.3 2+2+1

+ 4. 222531

1 3535.

545.

the proportion of granite-derived material in the soil is

greater.

1. 1.3.1 Rhus angustifolia-Berzelia lanuginosa Short

Closed Shrubland (D)

Campbell et al. (1981) structural equivalent: Low Closed Gra-

minoid Shrubland

Differential species: the differential species of this Community

are shared with the Benelia lanuginosa-Merxmuellera cincta

variant: Berzelia lanuginosa, Elegia asperiflora, Neesenbeckia

punctoria. It is distinguished from the variant by the lack of

differential species unique to the variant

Dominant species: Berzelia lanuginosa, Elegia asperiflora, Te-

traria fasciata

The Rhus angustifolia-Berzelia lanuginosa Shrubland

(Figure 5) is found in the lower central part of the study

area (altitude: 380—530 m) which has a north-easterly

aspect.

The soils on which this Community occurs are moist

but well drained. Fry (1987) classified these soils into

three forms and three series: Westleigh Form, Witsand

Series (We 21); Femwood Form, Warrington Series

(Fw 31) and Vilafontes Form, Hudley Series (Vf 11).

The parent material is sandstone. The Witsand and War-

rington Series are described as ‘hydromorphic’ soils and

the Hudley Series as a ‘pale’ soil which is moderately to

poorly drained.

A low stratum 0,5 m high with an average cover of

87% dominated by sedges, notably Tetraria fasciata is

found. Restios, grasses and other herbaceous species

contribute to this layer as well. Berzelia lanuginosa

emerges from the low stratum, occurring sparsely with

an average cover of 10% and not reaching more than 1

m in height.

Osmitopsis asteriscoides , a ubiquitous seep species, is

absent from this Community owing to drier soil condi-

tions. The presence or absence of this species may there-

fore be used in distinguishing this Community from the

Berzelia lanuginosa-Merxmuellera cincta Shrubland.

1.1. 3. 1.1 Berzelia lanuginosa-Merxmuellera cincta Tall

Closed Shrubland (C)

Campbell et al. (1981) structural equivalent: Mid-high Mid-dense

Shrubland

Differential species: Cunonia capensis, Laurentia arabidea, Merx-

muellera cincta, Platycaulos depauperatus

Dominant species: Berzelia lanuginosa, Merxmuellera cincta, Os-

mitopsis asteriscoides

The Berzelia lanuginosa-Merxmuellera cincta Tall

Closed Shrubland (Figure 6) is a variant of the Rhus an-

gustifolia-Berzelia lanuginosa Shrubland. It occurs in

conditions similar to those of the latter Community and

246

Bothalia 18,2 (1988)

the two Communities are found closely associated in the

lower central area of Swartboschkloof. It also occurs as a

more distinct unit along a drainage line on the slopes

above Sosyskloof stream.

The soils of the Fernwood Form, Warrington Series,

in the lower central area where the Berzelia lanuginosa-

Merxmuellera cincta Shrubland is found, have a greater

amount of accumulated organic material on the soil

surface on average (91%) than where the Rhus angusti-

folia- Berzelia lanuginosa Shrubland occurs. The diag-

nostic horizon of the soils of the Warrington Series con-

sists of acid, medium-grade, wet regie sand.

Structurally the Berzelia lanuginosa- Merxmuellera

cincta Shrubland differs from the typical community in

that the upper stratum is dominant. This stratum is 1-2

m high, has an average projected canopy cover of 60%

and is dominated by B. lanuginosa, M. cincta, Neesen-

beckia punctoria and Osmitopsis asteriscoides. M. cincta

and N. punctoria grow as large tussocks between which

B. lanuginosa and O. asteriscoides emerge. The lower

FIGURE 5. — Rhus angustifolia-

Berzelia lanuginosa Short '

Gosed Shrubland (D) in the

foreground.

stratum (<1 m) has an average projected canopy cover

of 45% and is dominated by Elegia asperi folia, Erica

hispidula and Platycaulos depauperatus .

Evidence suggests that the development of the Berze-

lia lanuginosa-Merxmuellera cincta Shrubland depends

largely on the presence of a high water-table.

Analysis of the Werger et al. (1972) data shows that

the community they described as the Berzelia lanuginosa

-Osmitopsis asteriscoides Community is the same as the

Berzelia lanuginosa-Merxmuellera cincta Shrubland de-

scribed here.

1 . 1 .3.2 Rhus angustifolia-Zantedeschia aethiopica High

Closed Shrubland (E)

Campbell et al. (1981) structural equivalent: Tall Gosed Proteoid

Shrubland

Differential species: the differential species of this Community

are shared with the Myrsine africana-Olea europaea subsp. afri-

cana Shrubland. They are: Blechnum australe, Maytenus acumi-

nata, Olea europaea susbp. africana and Zantedeschia aethiopica

FIGURE 6. — Berzelia lanuginosa

-Merxmuellera cincta Tall

Closed Shrubland. Note the

erect Osmitopsis asteriscoi-

des emergent through the

graminoid layer.

Bothalia 18,2 (1988)

247

Dominant species: Pteridium aquilinum. Protea neriifolia, Rhus

angustifolia

The Rhus angustifolia-Zantedeschia aethiopica High

Closed Shrubland (Figure 7) occurs in the mid-central

part of the study area below the contour path at altitudes

ranging from 450—530 m.

Three releves were situated in stands on soils of the

Champagne Form (Ch 1 1) and the other two in vegeta-

tion on soils of the Nomanci Form (No 10) and Magwa

Form (Ma 10) respectively (Fry 1987).

The tall form of the Rhus angustifolia-Zantedeschia

aethiopica Shrubland is found on Champagne Form soils.

The short stratum (<1 m) has the highest projected

canopy cover (93%) and is dominated by Pteridium

aquilinum and Z. aethiopica. R. angustifolia is emergent

from this stratum forming a shrub layer (1-2 m) with a

projected canopy cover of 22%. This form of the Com-

munity is clearly distinguishable on aerial photographs

and in the field, appearing as a physiognomic entity

separate from the other form.

The second form has a high stratum (2—4 m) with a

projected canopy cover of 70% . Protea neriifolia is the

dominant shrub in this stratum. It can also have a tall

stratum (1-2 m) which, if present, is usually dominated

by Brunia nodiflora and Cliff ortia cuneata. The lowest

stratum is a short (<1 m) closed layer (56% projected

canopy cover) with Pteridium aquilinum dominant. This

form of the Community occurs on soils of the Nomanci

and Magwa Forms (Fry 1987).

It may be argued that the ‘tali’ form of the community

is a separate community from the ‘high’ form, on the

basis of physiognomic appearance and structure. How-

ever, on the basis of floristic composition, a distinction

could not be drawn between them. These two structural

forms are therefore mapped as a single unit on the map

of plant communities.

1 .1 .3.3 Rhus angustifolia-Myrsine africana Short to

High Closed Shrubland

This shrubland consists of two variants, the Myrsine

africana-Olea europaea subsp. africana High Closed

Shrubland and the Myrsine africana-Cliffortia dentata

Tall Closed Shrubland. All the soils on which these Com-

munities occur, except Clovelly Form, Geelhout Series

(Cv 11), have granite material in the profile. The influ-

ence of granite is apparently greater on the former than

on the latter Community. In the soils of the Myrsine

africana-Qiffortia dentata Shrubland (except in the

Glenrosa Form), sandstone-derived materials are also

well represented.

The average altitudes at which these two Communi-

ties occur differ by over 160 m. The gradient of slopes

where the Myrsine africana-Cliffortia dentata Shrub-

land occurs average 38° which is almost twice as steep

as those on which the Myrsine africana-Olea europaea

subsp. africana Shrubland is found. The steeper slopes

are also cooler and moister. They have a north-easterly

to south-easterly aspect in contrast to the north-easterly

to north-westerly aspect of the area where the Myrsine

africana-Olea europaea subsp .africana Shrubland occurs.

Floristically, the two variants are related by having a

number of general (ubiquitous) species in common. More

specifically, these shrublands have three species, Protas-

paragus compactus, Psoralea monophylla and Tetraria

sylvatica which are exclusive to them.

1 . 1 .3.3. 1 Myrsine africana-Olea europaea subsp. africana

High Closed Shrubland (F)

Campbell et al. structural equivalent: Tall Closed Proteoid Shrub-

land

Differential species: the differential species of this Community

are shared with the Rhus angustifolia-Zantedeschia aethiopica,

Myrsine africana- Cliff ortia dentata and Ischyrolepis gaudichau-

diana-Myrsine africana Shrubland Communities. They are as

follows: Protasparagus compactus, Blechnum australe, Halleria

lucida, Hartogiella schinoides, Maytenus acuminata, Myrsine

africana, Olea europaea subsp. africana, Rapanea melanophloeos,

Rhus tomentosa, Tetraria sylvatica, Zantedeschia aethiopica

FIGURE 7. — Rhus angustifolia-

Zantedeschia aethiopica

High Closed Shrubland.

Note ‘tall’ form in the fore-

ground and the ‘high’ form

behind.

248

Dominant species: Cliffortia cuneata, Diospyros glabra, Olea

europaea subsp. africana, Protea neriifolia, Pteridium aquilinum,

Rhus angustifolia

The Myrsine africana-Olea europaea subsp. africana

High Closed Shrubland is found in the mid-central part

of the study area. It does not occur as a single unit but

rather as disjunct patches interspersed between other

communities.

The soils on which this Community is found are all

granite-derived and include the same three forms and

series as for the Rhus angustifolia- Zantedeschia aethi-

opica Shrubland: Champagne Form (Ch 11), Nomanci

Form (No 10) and Magwa Form (Ma 10).

The Myrsine africana-Olea europaea subsp. africana

Shrubland is characterized by a high stratum (2—6 m)

with an average projected canopy cover of 67%. This

stratum is dominated by O. europaea subsp. africana,

Protea neriifolia and Rhus angustifolia. A tall stratum

(1—2 m) with an average projected canopy cover of

25% is found in some stands. It is dominated by Clif-

fortia cuneata. A short stratum (<1 m) with an average

projected canopy cover of 24% and dominated by Pteri-

dium aquilinum is the typical understorey of this Com-

munity.

1.1. 3. 3. 2 Myrsine africana- Giffortia dentata Short

Closed Shrubland (G)

Campbell et al. (1981) structural equivalent: Low Closed Shrub-

land

Differential species: Giffortia dentata, Ficinia trichodes, Galium

mucroniferum, Pelargonium cf. tabiilare, Pentaschistis aristidoi-

des, Polyarrhena reflexa, Selago serrata

Dominant species: Aristea major, Giffortia cuneata, Pteridium

aquilinum

The Myrsine africana-Cliffortia dentata Short Closed

Shrubland is found on cool north-east to south-east-facing

slopes, in the south-west sector of Swartboschkloof. It

ranges in altitude from 680—700 m. The slopes are steep,

varying from 32°— 42°. This is much steeper than the

average gradient of the slopes in the catchment.

Four different soil series are found in the four releves

representing this Community: Hutton Form, Clansthal

Series (Hu 24); Glenrosa Form, Glenrosa Series (Gs 15);

Clovelly Form, Mossdale Series (Cv 14) and Clove lly

Form, Geelhout Series (Cv 1 1).

The dominant stratum of the Myrsine africana-

Cliffortia dentata Shrubland is a short layer (< 1 m) with

an average projected canopy cover of 72%. This stratum

is dominated by Aristea major, C. dentata. Erica sphae-

roidea and Pteridium aquilinum. In moist situations, C.

dentata forms extensive low mats. Above the short stra-

tum is a tall shrub layer with an average projected canopy

cover of 35%. It is dominated by A. major, Cannomois

virgata and Giffortia cuneata. C. dentata, Ficinia tricho-

des and Selago serrata are common to the Diospyros

glabra-Elegia capensis Shrubland and the Myrsine afri-

cana-Giffortia dentata Shrubland. This suggests some

affinity between these two Communities, possibly due

to the moist conditions in which they are found.

Bothalia 18,2 (1988)

1 . 1 .3.4 Rhus angustifolia-Ischyrolepis gaudichaudiana

High Closed Shrubland (I)

Campbell et al. (1981) structural equivalent: Tall Closed Pro-

teoid Shrubland

Differential species: this Community has no true differential

species. It is characterized on the basis of low occurrence or

absence of Hartogiella schinoides, Myrsine africana, Oxalis bi-

fida and Rhus tomentosa

Dominant species: Aristea major, Merxmuellera stricta, Protea

neriifolia, Pteridium aquilinum, Ischyrolepis gaudichaudiana

The Rhus angustifolia-Ischyrolepis gaudichaudiana

High Closed Shrubland in its typical form occurs inter-

spersed with the Rhus angustifolia-Myrsine africana

Shrubland over a large part of the mid-central zone of

the study area. It is found on a wide variety of soils:

Champagne Form (Ch 11), Magwa Form (Ma 10), Clovel-

ly Form (Cv 11, 14) and Glenrosa Form (Gs 15). At

elevations between 550 and 700 m, on the central mid-

slopes, it occurs on soils of Nomanci and Glenrosa Forms.

Protea neriifolia and P. repens dominate the high shrub

layer (2—5 m) which has an average projected canopy

cover of 34%. Following Edwards (1983), this is a mode-

rately closed canopy. Seven of the 18 releves sampled

lacked this stratum.

A tall stratum (1—2 m) occurs in all but one releve.

It has an average projected canopy cover of 25% and is

dominated by Brunia nodi flora, Cannomois virgata, Gif-

fortia cuneata, C. ruscifolia, Diospyros glabra, Halleria

elliptica and Leucadendron salignum. The short stratum

(<1 m) has the highest average projected cover (64%).

Dominant species in this stratum are: Aristea major,

Cymbopogon marginatus, Merxmuellera stricta, Pteri-

dium aquilinum, Ischyrolepis gaudichaudiana and Wat-

sonia pyramidata.

1 . 1 . 3 .4 . 1 Ischyrolepis gaudichaudiana-Myrsine africana

High Closed Shrubland (H)

Campbell et al. (1981) structural equivalent: Tall Closed Pro-

teoid Shrubland

Differential species: this Community has no true differential

species. It is distinguished from the Rhus angustifolia-Ischyro-

lepis gaudichaudiana Shrubland by the marked presence of

Hartogiella schinoides, Myrsine africana, Oxalis bifida and

Rhus tomentosa

Dominant species: Giffortia ruscifolia, Cymbopogon margina-

tus, Merxmuellera stricta. Protea neriifolia, P. nitida, Pteridium

aquilinum, Ischyrolepis gaudichaudiana

The Ischyrolepis gaudichaudiana-Myrsine africana

High Closed Shrubland (Figure 8) is found at altitudes

from 330-500 m in the lower to mid-central parts of

the study area, associated with the Rhus angustifolia-

Myrsine africana Shrubland. The soils on which it is

found are: Magwa Form (Ma 10), Clovelly Form (Cv

1 1), Oakleaf Form (Oa 34) and Glenrosa Form (Gs 15).

Three strata are distinguished in this Community. The

high stratum (2-5 m) is dominated by Protea nitida, P.

neriifolia and less commonly by Cliffortia ruscifolia and

P. repens. The average projected canopy cover for this

stratum is 44%. The tall (1-2 m) stratum which is do-

Bothalia 18,2 (1988)

249

FIGURE 8. — Ischyrolepis gaudi-

chaudiana-Myrsine africa-

na High Closed Shiubland.

Note the Protea nitida

shrubs, one of the domi-

nant shrubs in the tall to

high stratum of this com-

munity.

minated by C. cuneata, Diospyros glabra , Rhus angusti-

folia, Halleria elliptica and C. rusci folia, has an average

projected canopy cover of 22%. The short stratum (<1

m) has the highest average projected canopy cover (66% ).

It is dominated by Aristea major, Cymbopogon margi-

natus, Merxmuellera stricta, Montinia caryophyllacea,

Pteridium aquilinum and Ischyrolepis gaudichaudiana .

1.2 Diospyros glabra-Protea repens Transitional

Shrublands

The Diospyros glabra-Protea repens Shrublands

(Table 2) consist of those communities that are transi-

tional between the Erica hispidula- Diospyros glabra

Shrublands and the Erica hispidula- Ischyrolepis sieberi

Shrublands. There are two groups, the Protea repens-

Rhus angustifolia High Closed Shrubland and the Protea

repens-Nebelia paleacea High Closed Shrubland. The

first group has a typical community with two variants

and the second has two variants.

All these Communities, except for two releves on soils

derived mainly from granite, occur on sandstone-derived

soils. The Protea repens-Rhus angustifolia Shrublands

are more akin to the ‘pure’ communities found on soils

of granite and mixed (granite-sandstone) derivation. In

contrast, the Protea repens-Nebelia paleacea Shrublands

are allied to the sandstone-associated communities of the

Erica hispidula- Ischyrolepis sieberi Shrublands.

Table 2 indicates a fall-off in the occurrence of those

species with a granite or mixed soil affiliation as one pro-

ceeds from left to right through the transitional com-

munities. A sharp increase in the occurrence of sandstone-

favouring species is found in the Protea repens-Nebelia

paleacea Shrubland Communities compared with the

Protea repens-Rhus angustifolia Shrubland Communi-

ties.

Protea repens has been used in the binomial of the

major group because there is no other species consider-

ed suitable for this purpose (Table 2). It may be argued

that it is not a suitable species because it has a wide

ecological tolerance and is adversely affected by fire.

However, Protea repens is a conspicuous, easily identi-

fied shrub, which performs best in the transitional com-

munities. These attributes outweigh those not in its

favour.