FLORA

OF SOUTHERN AFRICA

VOLUME I

EDITED BY

L. E. CODD B. DE WINTER

AND

H. B. RYCROFT

Price R1.75 Overseas R2.20

Post Free

PUBLISHED IN THE REPUBLIC OF SOUTH AFRICA AND

PRINTED BY CAPE AND TRANSVAAL PRINTERS LIMITED

Digitized by the Internet Archive

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https://archive.org/details/floraofsoutherna01unse

FLORA OF SOUTHERN AFRICA

which deals with the territories of

THE REPUBLIC OF SOUTH AFRICA, BASUTOLAND,

SWAZILAND AND SOUTH WEST AFRICA

VOLUME I

Edited by

L. E. CODD and B. DE WINTER

Botanical Research Institute,

Department of Agricultural Technical Services

and

H. B. RYCROFT

National Botanic Gardens, Kirstenbosch,

Department of Education, Arts and Science

1966

Published in the Republic of South Africa and printed by Cape and Transvaal Printers Limited

THE TERRITORIES DEALT WITH IN THIS FLORA

baliniiJ 3i9JniiT

CONTENTS

Page

Introduction vii

Plan of Flora viii

Stangeriaceae by R. A. Dyer 1

Zamiaceae by R. A. Dyer and I. C. Verdoorn 3

Podocarpaceae by O. A. Leistner 34

Pinaceae by J. P. Jessop 42

Cupressaceae by J. A. Mart>h 43

Welwitschiaceae by I. C. Verdoorn 48

Cultivated Gymnosperms by R. J. Poynton 51

Typhaceae by J. G. Anderson 53

Helobiae by A. A. Obermeyer 56

Zosteraceae by A. A. Obermeyer 57

Potamogetonaceae by A. A. Obermeyer .... 60

Ruppiaceae by A. A. Obermeyer 70

Zanichelliaceae by A. A. Obermeyer 73

Najadaceae by A. A. Obermeyer 81

Aponogetonaceae by A. A. Obermeyer 85

Juncaginaceae by A. A. Obermeyer 92

Alismataceae by A. A. Obermeyer 96

Hydrocharitaceae by A. A. Obermeyer 100

Index 113

v

INTRODUCTION

THE second part of the Flora of Southern Africa to be published is Volume 1 of the planned

series, as set out on pp. viii and ix. The first part to appear, Vol. 26 in the series, was

published in 1963. No particular sequence of publication has been fixed and each volume is

sent to press as it is completed, irrespective of its place in the proposed plan of the whole work.

For convenience the families are arranged on the lines of the Engler system, largely because

the Flora is designed to be used in conjunction with the Genera of South African Flowering

Plants by E. P. Phillips. The delimitation of families does not necessarily follow the latter

work and modern trends may be accepted where such a procedure is considered justified.

The genera are numbered in the Flora on the basis of the system published by De Dalla Torre

and Harms in order to facilitate reference in Herbaria, such as those administered by the

Botanical Research Institute, which have adopted this system of arranging their material.

Vol. 1 includes the Gymnosperms, the Typhaceae and the early families of water plants

belonging to the Order Helobiae. The families in the Gymnosperms and Helobiae have been

delimited on the lines that are now more or less generally accepted and deviate, in certain

respects, from the treatment in the 2nd edition of Phillips’s Genera (1951). A key to the

families included in the Helobiae is, therefore, given on pp.

As in Vol. 26 (1963), the generally accepted abbreviations are used except in the following

cases which appear frequently and are therefore considerably condensed: —

F.C Flora Capensis

F.T.A Flora of Tropical Africa

F.T.E.A Flora of Tropical East Africa

F.Z Flora Zambesiaca

I.C.B.N International Code of Botanical Nomenclature

Phill., Gen The Genera of South African Flowering Plants by E. P.

Phillips, ed. 2 (1951)

Burtt Davy, FI. Transv. . . . Manual of the Flowering Plants and Ferns of the Trans-

vaal and Swaziland, Vol 1 (1926) and Vol. 2 (1932).

As before, the abbreviation “1. c.” is used for previously cited references, even though

“op. cit.” or “tom. cit”’ would in certain cases be more correct.

Magisterial districts are cited alphabetically in each Province and the intention is to follow

the latest delimitations of districts except where they are very small when a more general term

may be used, for example : —

Peninsula includes the Districts of Cape Town, Wynberg and Simonstown.

Reef includes the Districts of Roodepoort, Johannesburg, Germiston, Alberton, Kempton

Park, Benoni, Brakpan and Springs.

The spelling of the names of some localities has been brought into line with the findings of

the Committee on Standardisation of Place Names.

In the Index, synonyms are in italics and exotic species are signified by an asterisk*.

Although the name of Miss I. C. Verdoorn is not listed among the Editors of this work, it

is a pleasure to acknowledge the great debt owed to her for her constant stimulus in furthering

the progress of the work and for her careful scrutiny of the nomenclature and wording of the

entire text. Others who have assisted the Editors in many ways are Dr. R. A. Dyer, former

Chief of the Institute, Dr. D. J. B. Killick, Mrs. Obermeyer-Mauve and Mr. J. P. Jessop.

vii

PLAN OF CONTENTS OF VOLUMES

Stangeriaceae

Zamiaceae

Podocarpaceae

Pinaceae

Cupressaceae

Welwitschiaceae

Typhaceae

Zosteraceae

Potamogetonaceae

Ruppiaceae

Zanichelliaceae

Najadaceae

Aponogetonaceae

Juncaginaceae

Alismataceae

Hydrocharitaceae

Gramineae

Cyperaceae

Palmae

Araceae

Lemnaceae

Flagellariaceae

Restionaceae

Mayacaceae

Xyridaceae

Eriocaulaceae

Commelinaceae

Pontederiaceae

Juncaceae

Liliaceae

Flaemodoraceae

Amaryllidaceae

Velloziaceae

Dioscoreaceae

Iridaceae

Musaceae

Zingiberaceae

Burmanniaceae

Orchidaceae

Piperaceae

Salicaceae

Myricaceae

Ulmaceae

Moraceae

Urticaceae

Proteaceae

Vol. 1

Vol. 2

Vol. 3

Vol. 4

Vol. 5

Vol. 6

Vol. 7

Vol. 8

Vol. 9

Vol. 10

Loranthaceae

Santalaceae

Grubbiaceae

Olacaceae

Balanophoraceae

Aristolochiaceae

Rafflesiaceae

Hydnoraceae

Polygonaceae

Chenopodiaceae

Amaranthaceae

Nyctaginaceae

Vol. 11

Phytolaccaceae

Aizoaceae

Vol. 12

Portulacaceae

Basellaceae

Caryophyllaceae

Nymphaeaceae

Ceratophyllaceae

Ranunculaceae

Menispermaceae

Annonaceae

Monimiaceae

Lauraceae

Flernandiaceae

Papaveraceae

Fumariaceae

Vol. 13

Cruciferae

Capparidaceae

Resedaceae

Moringaceae

Droseraceae

Podostemaceae

Flydrostachyaceae

Vol. 14

Crassulaceae

Vol. 15

Saxifragaceae

Pittosporaceae

Cunoniaceae

Myrothamnaceae

Bruniaceae

Hamamelidaceae

Rosaceae

Connaraceae

Vol. 16

Leguminosae

Vol. 17

Geraniaceae

Oxalidaceae

viii

PLAN OF CONTENTS OF VOLUMES

Vol. 18

Linaceae

Erythroxylaceae

Zygophyllaceae

Rutaceae

Simarubaceae

Buseraceae

Meliaceae

Malpighiaceae

Vol. 19

Polygalaceae

Dichapetalaceae

Euphorbiaceae

Callitrichaceae

Buxaceae

Anacardiaceae

Aquifoliaceae

Vol. 20

Celastraceae

Hippocrateaceae

Icacinaceae

Sapindaceae

Melianthaceae

Balsaminaceae

Rhamnaceae

Heteropyxidaceae

Vitaceae

Vol. 21

Tiliaceae

Malvaceae

Bombacaceae

Sterculiaceae

Vol. 22

Ochnaceae

Guttiferae

Elatinaceae

Frankeniaceae

Tamaricaceae

Canellaceae

Violaceae

Flacourtiaceae

Turneraceae

Passifloraceae

Achariaceae

Loasaceae

Begoniaceae

Cactaceae

Vol. 23

Geissolomaceae

Penaeaceae

Oliniaceae

Thymelaeaceae

Lythraceae

Lecythidaceae

Vol. 24

Rhizophoraceae

Combretaceae

Myrtaceae

Melastomataceae

Onagraceae

Hydrocaryaceae

Halorrhagidaceae

Araliaceae

Umbelliferae

Cornaceae

Vol. 25

Ericaceae

Vol. 26

Myrsinaceae

Primulaceae

Plumbaginaceae

Sapotaceae

Ebenaceae

Oleaceae

Salvadoraceae

Loganiaceae

Gentianaceae

Apocynaceae

Vol. 27

Asclepiadaceae

Vol. 28

Convolvulaceae

Hydrophyllaceae

Boraginaceae

Verbenaceae

Labiatae

Solanaceae

Vol. 29

Scrophulariaceae

Vol. 30

Bignoniaceae

Pedaliaceae

Martyniaceae

Orobanchaceae

Gesneriaceae

Lentibulariaceae

Acanthaceae

Myoporaceae

Vol. 31

Plantaginaceae

Rubiaceae

Valerianaceae

Dipsacaceae

Cucurbitaceae

Vol. 32

Campanulaceae

Goodeniaceae

Vol. 33

Compositae

IX

STANGERIACEAE

by R. A. Dyer

Plants dioecious with thick roots merging into simple or branched subterranean stems.

Stem tuberous, with leaves from the apex of each stem or branch, alternating with series of

small scale-like bracts. Leaves pinnate, 1-4 from each stem, somewhat fern-like, petiolate,

deciduous; leaflets in several to many opposite or subopposite pairs, with a conspicuous

midrib and dichotomously forked lateral veins. Cones with scales (sporophylls) arranged

spirally, tightly imbricate upwards. Male scales with the pollen cells stipitate on the lower

surface towards the base; pollen ellipsoidal. Female scales stalked with two ovules, one on

either side and above the pedicel with the micropyle directed towards the cone axis; scale-head

or bulla shield-like with incurved lateral lobes. Seeds with fleshy outer layer tightly adhering

to the hard inner shell encasing the copious endosperm; cotyledons 2.

A monotypic family endemic in the eastern coastal region of South Africa.

In the past most workers have referred all living cycads to a single family. L. A. S. Johnson, in the Pro-

ceedings of the Linnaean Society of New South Wales 84 : 64 (1960), restricts the family Cycadaceae to the

single genus Cycas. He creates the new family Stangeriaceae and resuscitates the family Zamiaceae for the

remaining genera. Johnson’s classification is supported by Kew and is followed here in the interests of a

uniform nomenclature.

2

STANGERIA

Stangeria T. Moore in Hook., Journ. Bot. 5 : 228 (1853); J. Smith in Hook., ibid 6 : 88 (1854);

Marloth, FI. S. Afr. 1 : 97, t. 14, t. 63 (1913); Pilg. in Pflanzenr. 15 : 74 (1926); Schuster in

Pflanzenr. 4, 1 : 103 (1932); Hutch. & Rattr. in F. C. 5, 2 (Suppl.) : 25 (1933); Phill., Gen. ed.

2 : 49 (1951); R. A. Dyer in Bothalia 8 : 429 (1965).

Description as for family.

A monotypic genus on the eastern border of South Africa; named in honour of Dr. William Stanger,

who became Surveyor General of Natal.

Stangeria eriopus ( Kunze ) Nash in Journ.

New York Bot. Gard. 10 : 164, pi. 62 (1909);

Hutch. & Rattr. in F.C. 5, 2 (Suppl.): 26

(1933); R. A. Dyer in Bothalia 8 : 429 (1965).

Type: eastern Cape, Drege.

Lomaria eriopus Kunze in Linnaea 13 : 152

(1839); 18 : 116 (1844). L. coriacea sensu Kunze

in Linnaea 13 : 152 (1839).

Stangeria paradoxa T. Moore in Hook., Journ.

Bot. 5 : 228 (1853); Hook, in Bot. Mag. t.5121

(1859); Schuster in Pflanzenr. 4, 1 : 103; fig. 15A-K

(1932). Type: Natal, Stanger s.n. — var. katzeri

(Regel) Marloth, FI. S. Afr. 1 : 97, fig. 63 (1913);

Schuster l.c. 105, fig. 15B, t.3 (1932). - — var. schizodon

(Bull.) Marloth, l.c. pi. 14 (1913). — forma schizodon

(Bull.) Schuster, l.c. 105, fig. 15C, t.3 (1932).

S. schizodon Bull., Cat. 1872 : 8 (1872). Type;

Hort. S. katzeri Regel, Gartenfl. 23 : 163, t.798

(1874). Type: Hort. S. zeyheri Stoneman, Plants

and their Ways in S. Afr., fig. 214 (1915), nom.

subnud.

Perennial herb with variously carrot-

shaped main root or roots continuous with

the main stem. Stem subterranean, tuberous,

turnip-shaped, up to 10 cm diam., branched

or unbranched, with ovate, early deciduous,

woolly scales at the apex. Leaves pinnate, 1-4

from the apex of each stem or branch, ex-

tremely variable in size (according to ecologi-

cal conditions), 25 cm-2 m tall, petiolate;

petiole more or less equal to the length of the

divided blade, grooved on upper side; leaflets

in 5-20 pairs, opposite or subopposite, the

lowermost shortly stalked, the uppermost

2

Stangeriaceae

Fig. 1. — Stangeria eriopus, a, typical leaf from open veld; b, leaflet from forest locality; c, portion of leaflet

showing veins; d, margin of serrulate leaflet; much reduced except c and d (taken from F.C. 5, 2 (Suppl.)

1933.

Stangeriaceae

3

connate and decurrent on the rhachis on the

lower side, entire or serrulate to irregularly

incised-lobate, elongate-lanceolate, acutely

acuminate to rounded at apex, up to 40 cm

long and 6 cm broad, glabrous. Cones

solitary, silvery pubescent becoming brown

tomentose, pedunculate. Male cones sub-

cylindric, slender, tapered to the apex, 10-15

cm long and up to 3 cm diam., with numerous

spirals of scales closely imbricate upwards;

scales glabrescent, broadly triangular or

rhomboid; bulla face curved upwards with

sharp teeth on margin. Female cones elliptic-

ovoid, up to 18 cm long, 8 cm diam.; scales

stalked; bulla with slightly convex peltate

face, imbricate upward, irregularly toothed

on margin. Seed red, up to 3-5 cm long, 2-5

cm diam., angled by compression, with fleshy

covering adhering firmly to the thin hard

inner coat encasing the copious endosperm.

Fig. 1.

In coastal grassveld and forest from Bathurst

District in the eastern Cape to northern Natal

(Zululand) near Kosi Bay.

Cape.- — Bathurst: MacOwan 2000; Dyer 4836.

East London: Galpin 7366; Dyer 4528. Kentani:

Pegler 262; 1247; Flanagan 1118; Dyer 4514;

4516. Port St. Johns: Galpin 2849.

Natal. — Port Shepstone: Mogg 13906; McClean

352. Umzinto: FUlliard 3121. Pinetown: Strey

3852; Bruce 151. Mtunzini: Codd & Dyer 2818.

Eshowe: Acocks 12970. Ingwavuma: Ward 3717;

Vahrmeijer & Tolken 995.

This plant was originally confused with a fern as

the synonymy shows. The tall graceful leaves pro-

duced by plants within forests are very different

from the relatively small hard ones characteristic

of plants in open grassveld. It does not seem possible

to draw a clear line of distinction between the forms,

which fact makes it unsatisfactory to distinguish

taxa of varietal rank. The tuberous rootstock is

regarded as all underground stem by some writers

but the basal portion is a tuberous root which

develops rapidly in the seedling stage. The transi-

tional area between stem and root is not readily

detected. The stem portion may occassionally

branch into nearly a dozen heads, each of which

may produce a cone at the same time.

ZAMIACEAE

by R. A. Dyer and I. C. Verdoorn

Plants dioecious, with subterranean or aerial stems. Stem protected by persistent leaf-

bases, fibrous round the circumference and with central fibrous pith, branched or unbranched

from the base, rarely above, with a few to many leaves produced from the apex of the stems

and alternating with series of ovate to lanceolate woolly bracts. Leaves pinnate, palm-like.

Cones with male and female scales (sporophylls) spirally arranged, 1-several produced at the

apex of the stems. Male scales with the pollen cells produced densely or in groups on the

basal part of the lower surface. Female scales very closely packed and persisting on the cones

for several months, pedicellate with an enlarged peltate head (bulla); bulla with incurved

lateral lobes and from the inner surface, one on either side above the pedicel, are produced

2 ovules with the micropyle directed towards the axis of the cone. Seeds with fleshy covering,

oblong in outline, angled by compression, with a thin hard globose or ovoid inner shell

encasing the copious endosperm; cotyledons 2.

Restored to family rank by Johnson in Proc. Linn. Soc. New S. Wales 84(1960). See note under Stangeria-

ceae.

Genera 8, scattered mainly in the southern hemisphere in tropical, subtropical and temperate America,

Australia and Africa; only one, Encephalartos, occurs in Southern Africa.

4

Ad axial view of

female cone scale

Zamiaceae

ALeiOA x. t>. WERWC

Fig. 2 — Main features of Encephalartos female cone scales.

Zamiaceae

5

5 ENCEPHALARTOS

Encephalartos Lehm. in Pugill. 6 : 3, t.1-5 (1834); De Yriese in Tydsch. Nat. Gesch. 4 : 411

(1838); Miq., Monogr. Cycad. 39 (1842); De Cycad. Loddig. (1842); Linnaea 17 : 709 (1843);

A. DC. in DC., Prod. 16, 2 : 530 (1868); Ed. Gard. Chron. 401 (1875); Pearson in Trans.

S. Afr. Phil. Soc. 16 : 345 (1906); Marloth, FI. S. Afr. 1 : 99(1913); Pearson in Joum. Bot. Soc.

S. Afr. 2 : 7 (1916); Chamberlain, The Living Cycads 45 (1919); Pilg. in Pflanzenfam. 13 : 78

(1926); Burtt Davy in FI. Transv. 1 : 39; 98 (1926); Schuster in Pflanzenr. 4, 1 : 105 (1932);

Hutch. & Rattr. in F.C. 5, 2 (Suppl.) : 28 (1933); M. R. Henderson in Journ. Bot. Soc. S. Afr.

29 : 6 (1943); Journ. S. Afr. Bot. 11:5 (1945); Phill., Gen. ed. 2 : 49 (1951); Verdoom in

Afr. Wild Life 5 : 153 (1951); Trees in S. Afr. 8 : 3 (1956); Melville in Kew Bull. 1957 : 237

(1957); Thieret in Economic Bot. 12 : 3 (1958), partly; Verdoorn in Trees in S. Afr. 14 : 50

(1962); Whiting in Economic Bot. 17 : 271 (1963), partly; Third Conference on the Tox.

Cycads, Federation Proceed. 23 : No. 6, pt. 1 (1964), partly; Dyer in Bothalia 8 : 432 (1965).

Plants dioecious with main root or roots tuberous. Stem unbranched or branched from

the base, rarely branched above and then largely due to injury, subterranean or aerial,

occasionally up to 10 m tall, covered by alternating series of woolly bracts and persistent

swollen truncated leaf bases (pulvini). Leaves arising from the apex of the stems, spirally

arranged, persisting 2-several years, petiolate, pinnate with a straight or curved rhachis;

leaflets pungent-pointed, entire, toothed or lobed on one or both margins, reduced in size

towards base of rhachis, sometimes to prickles, with longitudinal parallel venation and no

midrib; leaflets of seedlings of all species with several teeth round apex. Cones 1-5, sometimes

more, from near or round apex of stem, glabrous to densely woolly, pedunculate; peduncle

of male cones somewhat longer than those of the female cones in the same species. Male cones

subcylindric; scales densely packed in many spiral rows, terminating in a sterile truncate apex

or beak; pollen cells in dense clusters on the basal portion of lower surface. Female cones

cylindric to oval, broader than the male; scales tightly packed, stalked, bearing 2 collateral

ovules arising from the inner surface and lying along the upper side of the stalk, with an

enlarged peltate head (bulla); bulla face flat or somewhat protruding and with lateral ridges

extending into the incurved lateral lobes, having in the main an upper, lower and terminal

facet. Seed red, yellow, amber or brownish with age, fleshy, oblong, angled by compression,

with apical fleshy end variable in length, with a thin hard globose or oblong-ovoid inner shell

encasing the copious endosperm, having a long maturing period on the cone whether fertile

or not. Fig. 2.

Widely distributed on the African Continent but absent from South West Africa, western Cape, O.F.S.

and Basutoland; 35 species are recognised, 26 of which occur in Southern Africa. The Latin name is derived

from the common name Bread Tree or Broodboom, referring to the making of bread by the Hottentots

and Bantu from the starchy pith within the trunks.

Cycas revoluta Thunb. from the East, which is found occasionally in cultivation in South Africa, and

C. thouarsii R.Br., which occurs naturally along the Mozambique coast, could be mistaken for species of

Encephalartos. They are readily distinguished by a mid-rib in the leaflets and by the branched female

inflorescence which is not cone-like.

In citing literature, where there is no reason to disagree with the references given by Hutchinson and Rat-

tray in FI. Cap. 5, 2 (Suppl.) 1933, these have frequently been omitted.

Leaflets 2-4 mm broad, 8-14 cm long, linear, entire with revolute margins, dark green; crown of stem

brown woolly; cones 2-5 together, densely woolly; male cone scales on stipe 1-1 • 5 cm long 1. ghellinckii

Leaflets 4 mm or more broad, margins usually thickened and sometimes recurved but not revolute:

Leaflets towards the base of the petiole deflexed in relation to the rhachis, the basal leaflets reduced to

a short or long series of prickles, those about the middle of the rhachis spreading at right angles

to the axis and recurving, 14-21 cm long, 8-13 mm broad, linear, attenuate, 13-21-veined on the

under surface and glaucous (silvery or bluish) on under surface when young, stems branched from

the base, up to 2 m long, rarely longer; face of male cone scales moderately greyish pubescent

7. inopinus

6

Zamiaceae

Leaflets spreading at right angles or ascending in relation to the axis of the rhachis (none deflexed),

glaucous or green:

Leaflets 4-8 mm broad, linear, entire, glaucous or somewhat silvery when young, sometimes

yellowish with age, rarely reduced to more than one prickle at base, venation conspicuous on

under surface ; cones densely woolly or only moderately so in E. laevifolius, seeds light brown

or brown-madder to ochre-yellow:

Cones with dense woolly covering:

Stems well developed above ground:

Stem crown conspicuously and rather persistently brown woolly; trunk 35-60 cm diam., up

to 3 m or more tall; cones 3-several together; median leaflets 10-17 cm long and about

7 mm broad, 7-9-nerved on under surface 3. friderici-guilielmi

Stem crown not conspicuously brown-woolly, trunk rarely up to 30 cm diam. and not often

up to 2 m tall :

Median leaflets 9-13 cm long, 4-5 mm broad, 5-6-nerved on under surface; margin

somewhat recurved; cones often single 2. cycadifolius

Median leaflets 10-13 cm long, 6-8 mm broad, finely 10-14-nerved on under surface,

flat; cones 1-4 together 4. lanatus

Stems rarely up to 20 cm above ground, forming dwarf branched clumps; cones single; median

leaflets 9-13 cm long, 4-6 mm broad, finely about 9-nerved on under surface 5. humilis

Cones with short greyish tomentum, not conspicuously woolly; median leaflets up to about

12 cm long, 5-7 mm broad, 10-12-nerved on under surface; trunk 1-3 m tall, rarely more

than 30 cm diam 6. laevifolius

Leaflets 1 cm or more broad, if less than 1 cm broad then bright green not glaucous; margins

entire or toothed ; cones glabrous or apical part of scale face with reddish or brownish hairs :

tStems usually well developed above ground, if subterranean then leaves glaucous as in E. horridus:

Leaves glaucous with bluish bloom when young and persisting so for several months at least,

contrasting with the old leaves:

Leaves positively glaucous when young, leaflets rarely reduced to more than one prickle

at base, median leaflets usually less than 2-5 cm broad:

Median leaflets mainly entire or occasionally with 1 or 2 teeth on lower margin:

Leaves slightly or markedly incurved near the tip; female cone-scales with slightly

rugose face 8. eugene-maraisii

Leaves slightly or markedly recurved towards tip :

Female cone-scales with irregularly warty face, the pulvinus of petiole often partly

hidden by stem bracts, leaves often pilose when young, leaflets usually over-

lapping above middle of rhachis 9. princeps

Female cone scales with smooth or slightly rugose face, pulvinus of petiole exposed

and prominent, with brown collar, leaves glabrous, leaflets often not over-

lapping 10. lehmannii

Median leaflets, some or most, with 1-3 spine-tipped lobes on the lower margin, glabrous:

Stems mainly above ground ; median leaflets 1 • 5-2 • 5 cm broad, some or all with 1 or 2

spine-tipped lobes on lower margin, and lobes sometimes twisted out of the plane

of the leaflet; female cone scales yellowish-green, rugose on face with the lateral

incurved lobes 2-3 cm long 11. trispinosus

Stems mainly subterranean, occasionally up to 45 cm tall, rarely taller, median leaflets

rarely less than 2-5 cm broad, deeply 1- or 2-lobed on lower margin and lobes

twisted out of plane of the leaflets, female cone scales blackish or reddish-brown,

ridged towards terminal facet but otherwise with a fairly smooth face, with the

lateral incurved lobes about 1 cm long 12. horridus

Leaves light or medium green, faintly glaucous; median leaflets usually more than 2-5 cm

broad, lower margin with 3 or occasionally 4 spine-tipped lobes 13. arenarius

Leaves light or dark green, with or without prickles or lobes along one or both margins, not

glaucous except sometimes in young stage of E. longifolius and E. arenarius:

Lower margin of median leaflets with 2-4 triangular pungent lobes, upper margin without

lobes or teeth:

Leaves medium green, faintly glaucous, glabrous; leaflets not prominently veined on under

surface 13. arenarius

Leaves dark green, finely pubescent when young, rigid, leaflets prominently veined on

undersurface 14. Iatifrons

Lower and upper margin of median leaflets entire or with one or more prickles or teeth:

Leaflets reduced in size towards base of rhachis but not to more than one or exceptionally

2 prickles:

Zamiaceae

7

Leaves 1-1-5 m long, suberect-spreading and recurved or arcuate in the upper half;

leaflets dark green often with bluish lustre and sometimes glaucous when young,

finely pubescent when young, median ones mainly oblong-lanceolate, up to 20 cm

long and 4 cm broad, generally overlapping on upper half of rhachis, margins not or

sparingly toothed 15. longifolius

Leaves 1—1 - 5 m long, nearly straight; leaflets bright green, median ones oblong-

lanceolate to lanceolate up to 15 cm long and 2-5 cm broad, rarely overlapping,

variably toothed on margins; incurved lateral lobes of female cone scales lacerate-

winged, about 3 cm long and extending nearly to axis 16. altensteinii

Leaflets reduced to several or many prickles towards base of rhachis:

Leaflets with a single terminal spine or point, broadest below or occasionally near

middle:

Leaflets often showing veins on lower surface but not in regular raised formation:

Leaflets spreading and straight from the rhachis, stems often branched from

base, rarely up to 7 m tall:

Leaves curved, bow-shaped giving an umbrella-like canopy; median leaflets

dark green, more or less ovate-lanceolate, up to 20 cm long and 5 cm

broad 17. woodii

Leaves spreading, straight or slightly curved in the upper third:

Median leaflets lanceolate to ovate-lanceolate, 16-23 cm long, 2 -5-4 -5 cm

broad; face of female cone-scales with prominent blunt rugosities;

incurved lateral lobes toothed or laciniate, 2-3 cm long 18. natalensis

Median leaflets mainly narrowly lanceolate, 12-17 cm long, 1-2-2 -2 cm

broad; face of female cone-scales without prominent excrescences;

incurved lateral lobes, laciniate, about 1 -8 cm long 19. lebomboensis

Leaflets spreading-recurved from rhachis, lanceolate to ovate-lanceolate, some-

times falcate, 10-20 cm long, 2-3-5 cm broad; stems rarely branched, many

over 6 m tall 20. transvenosus

Leaflets with about 20 regular conspicuously raised longitudinal veins on lower

surface, median leaflets linear-lanceolate, straight or somewhat falcate, 15-25 cm

long, 2-3 cm broad, rarely more 21. paucidentatus

Leaflets with 3-5 almost equal pungent lobes at the end, usually 3-4 cm broad, broadest

above or sometimes near middle, slightly narrowed to the base; cones salmon or

shrimp-pink to scarlet 22. ferox

tfStems dwarf, subterranean or rarely up to 20 cm exposed above ground, unbranched or some-

times branched; leaflets light or dark green, not glaucous, linear, or linear-lanceolate, with or

without marginal teeth:

Leaflets reduced to many prickles towards base of petiole, median ones averaging 15-20 cm

long, usually with one or two prickles on both margins; female cone scales with the

terminal facet extending almost to the basal margin of the bulla face or overlapping slightly

the scale below; male cones 6-5-12 cm diam 23. villosus

Leaflets reduced to one or two prickles at the base or sometimes none:

Leaflets, some or nearly all with one or two prickles on the margins, average length 12-14 cm,

occasionally up to 30 cm, 8-15 mm broad, female cone scales with terminal facet

generally slightly below central, rarely overlapping scale below, male cones 6-8 cm

diam 24. umbeluziensis

Leaflets (except on juvenile plants) entire or with 1-3 small teeth on lower margin, average

length 6-8 cm long, 8-11 mm broad:

Female cone scales with terminal facet on the face indistinct and with the lower margin

as low as or slightly overlapping the scale below; male cones less than 8 cm diam.;

median leaflets usually toothed on lower margin (Natal) 25. ngoyanus

Female cone scales with terminal facet on the face distinct, not extending to basal margin;

male cones 6-11 cm diam.; median leaflets usually without prickles on lower margin

(Cape) 26. caffer

1. Encephalartos ghellinckii Lem., lllustr.

Hort. 14 Miscell. 80 (1867); lllustr. Hort.

15, pi. 567 (1868); Hutch. & Rattr. in F.C. 5,

2 (Suppl.): 43 (1933); M.R. Henderson in

Journ. S. Afr. Bot. 11 : 57 (1945); R. A. Dyer

in Bothalia 8 ; 437 (1965). Type: Hort. Ver-

schaffelt, no specimen traced, but tab. in

lllustr. Hort. 15, pi. 567 (1868) adequate;

probably originated from Natal, see R. A.

Dyer in Journ. S. Afr. Bot. 31 : 120 (1965),

Zamia ghellinckii Hort. ex Lem., l.c.

Encephalartos gracilis G. Henderson, 111. Diet.

Gard. Suppl. 341 (1900), nomen.

Plant unbranched or branched from the

base; stems up to about 3 m tall, rarely up to

Zamiaceae

3-5 m and then usually leaning; 30-40 cm

diam., with brown woolly crown. Leaves up

to 1 m long, including petiole 25 cm long, at

first densely greyish tomentose, glabrescent

except for brown silky-tomentose pulvinus;

rhachis stiff, slender, straight or spreading-

upcurved and sometimes somewhat twisted,

yellowish; leaflets spreading-reflexed, dense

above middle of rhachis, more spaced and re-

duced in size towards its base, with thickened

entire revolute margins and generally incon-

spicuous venation which may be more evident

on upper surface than lower; median leaflets

8-14 cm long, 2-4 mm broad, pungent-tipped.

Cones 2-5 together, on short stout peduncles,

densely covered with matted brownish tomen-

tum. Male cone cylindric, curved, about 25

cm long, 6-7 cm diam., narrowed slightly to

both ends, median scales thin, broadly ob-

cordate, 2-2-5 cm long, 3-3-5 cm broad, on

slender stalk (stipe) 1-1 • 5 cm long; bulla face

2 cm broad and 1 cm thick vertically, very

little projecting, with acute lateral wings, up-

per and lower facets rounded or slightly

humped, terminal facet concave, about 7 mm

broad and 3 mm wide vertically. Female cone

barrel-shaped, up to about 22 cm long and

14 cm diam. ; median scales about 4 cm long,

4 • 2-6 cm broad and 2 • 5-3 cm thick vertically,

with short recurved lateral lobes; bulla face

woolly, only slightly projecting, nearly flat.

Seed golden-yellow, about 2 • 5 cm long and as

broad or broader.

On the southern foothills of the Drakensberg in the

eastern Cape and from 2,000 ft altitude in southern

Natal to higher altitudes northwards along the

mountain range as far as Mont-aux-Sources;

occasionally in fair abundance. Map. 1.

Cape. — Flagstaff: McLoughlin 513a; Verdoorn &

Christian 712; 712a. Tabankulu: Acocks 12259. Mt.

Ayliff: Smith 6833.

Natal. — Port Shepstone: Strey 5878. Umzinto:

Wood 13055. Richmond: Hilliard 3145. Estcourt:

Symons 498; Dyer 4771. Bergville: Hutchinson,

Forbes & Verdoorn 48; Galpin 9648; Dohse 72;

Dyer & Codd 4784.

This species is readily distinguished from all others

of the genus by its narrow leaflets with recurved

margins. In leaf characters it resembles species of

Cycas in the family Cycadaceae, which differ in

having a midrib in the leaflets. The leaflets of

specimens from the Drakensberg seem on the

average to be slightly broader and with less recurved

margins than those farther south from lower alti-

tudes toward the coast.

2. Encephalartos cycadifolius ( Jacq .)

Lehm., Pugill. 6 : 13 (1834); Miq., Monogr.

Map 1. — Distribution map of Encephalartos

ghellinckii, E. cycadifolius, E. friderici-guilielmi,

E. lanatus, E. humilis, E. laevifolius and E.

inopinus.

Cycad. 43 (1842); Schuster in Pflanzenr. 4,

1 : 108 (1932), in part; Hutch. & Rattr. in

F.C. 5, 2 (Suppl.): 42 (1933), for the lesser

part, excluding citations; M.R. Henderson in

Journ. S. Afr. Bot. 1 1 : 50 (1945), for the lesser

part, excluding most citations; R. A. Dyer in

Bothalia 8 : 439 (1965). Type: Cape, without

locality, the illustration in Jacq., Fragm. 27,

t.26 (1801), from a plant cultivated in Vienna,

leg. Scholl', see R. A. Dyer in Journ. S. Afr.

Bot. 31 : 116 (1965).

Zamia cycadifolia Jacq., Fragm. 27, t.26 (1801);

(it is doubtful if t.25 is from the same species).

Encephalartos eximius Verdoorn in Bothalia

6 : 426 (1954). Type: Cape, Bedford, Story 4021

(PRE, holo.).

Plants branching from the base; stems

up to about 60 cm tall, rarely 90 cm and about

25 cm diam.; bracts about 3 cm long and

2 ■ 5 cm broad at base, tomentose on outer sur-

face. Leaves about 60-90 cm long including

petiole 10-20 cm long, woolly and with some

long straight hairs when young, glabrescent

except for densely greyish tomentose pulvinus ;

rhachis fairly stiff, suberect sometimes slightly

twisted; leaflets somewhat glaucous when

young, pungent-tipped, upper ones spreading

from the rhachis horizontally or forming a

wide V, not or only slightly overlapping, re-

duced gradually in size to base and apex of

Zamiaceae

9

Fig. 3. — Encephalartos cycadifolius, a, typical leaf reduced; b, tip of leaflet, enlarged (Dyer 5333).

10

Zamiaceae

rhachis; median leaflets 9-13 cm long, 4—5

mm broad, entire, with thickened slightly re-

curved margins and with 5-6 prominent

sparsely pilose nerves on under surface. Cones

1 or 2 from a stem on short fairly stout pe-

duncles subtended by narrow dorsally tomen-

tose bracts 2-7 cm long, densely tawny or

off-white tomentose. Male cones subcylindric,

13-22 cm long, 4-5-7 cm diam., narrowing

slightly to the base and rounded apex;

median scales about 2 cm long, 1-8 cm

broad, with lateral ridges; bulla face (with

tomentum) 1-2 cm broad and 5-8 mm

wide vertically, but only about 2 mm thick

vertically without tomentum. Female cones

more or less oblong in outline with rounded

apex, 20-30 cm long, 16-18 cm diam. ; median

scales 4-4-5 cm long and 4 cm broad; bulla

about 3-4 cm broad and 1 • 5-2 cm thick verti-

cally, with incurved lateral lobes up to 1 cm

long, yellowish, with acute lateral ridges,

upper and lower facets more or less rounded;

terminal facet about 2 cm broad and 1 • 25-1 • 5

cm wide vertically. Seeds pale orange-yellow

through salmon-suffused orange to amber-

brown, about 3 cm long and 1 • 75-2 cm broad.

Fig. 3.

Restricted to a few areas on the Winterberg in

the districts of Bedford and Cradock. Map 1.

Cape.— Bedford : Story 4013; 4017; 4018;

4019; 4021; 4022; 4023; 4548; Aver 5331; 5333.

Cradock: Acocks (photo); Mar loth 2151.

Specimens of E. friderici-guilielmi from Queens-

town and Cathcart districts have been confused with

this species at various times but the former is more

robust and has the leaflets more densely spaced.

The problems are discussed in the Journ. S.Afr.

Bot. 31 : 116 (1955).

As regards the date of publication of Zamia

cycadifolia Jacq., Z. horrida Jacq. and Z. longifolia

Jacq., Mr. A. A. Bullock of Kew has supplied the

following information: “The Fragmenta seem to

have been published in fascicles and the accepted

dates are as follows: The preface, p. 2, indicates

publication of fascicle one, some time after March

1800 and Konig & Sims, Ann. Bot. 1 : 64, give 1801

for fascicles 1 & 2, pp. 1-34, tt. 1—37. Other fascicles

are indicated in the text”.

3. Encephalartos friderici-guilielmi Lehm.,

Pugill. 6 ; 8, t.l, 2 & 3 (1834); Miq., Monogr.

Cycad. 44 (1842); R. A. Dyer in Journ. S. Afr.

Bot. 31:117 (1965); Bothalia 8 : 442 (1965).

Type: Cape, eastern area, Ecklon & Zeyher.

E. acanthus Mast, in Gard. Chron. 1 1 : 810 (1878);

Thiselton-Dyer in Gard. Chron. 11 : 11 (1879).

Type: Hort. Bull., Cape, probably Queenstown.

E. cycadifolius sensu Auct., Schuster in Pflanzenr.

4. 1 : 108 (1932) partly; Hutch. & Rattr. in F.C. 5,

2 (Suppl.): 42 (1932) partly; M. R. Henderson in

Journ. S. Afr. Bot. 11 : 50 (1945) partly, as to

Queenstown and Cathcart specimens etc.

Plant unbranched or branched from the

base ; stems stout, up to about 4 m tall, 35-60

cm diam., eventually leaning or procumbent

with age, with open densely brown woolly

crown; interseriate bracts linear-lanceolate,,

up to 14 cm long, densely woolly on outer

surface. Leaves 1-1 • 2 mlongincluding petiole

about 30 cm lcng, undersurface of rhachis and

margins of leaflets loosely whitish woolly

when young, glabrescent, with pulvinus per-

sistently woolly; rhachis fairly stiff, straight

or recurved, often spreading lower than crown

when stem has cones; leaflets somewhat

glaucous when young, closely spaced and

overlapping in the upper half of leaf and in V

disposition, linear, shortly narrowed to an

amber, dark brown or black pungent apex,

with entire thickened margins, and with the

basal ones reduced in size, but not to prickles;

median leaflets 10-17 cm long and about 7

mm broad, with 7-9 prominent nerves on the

lower surface. Cones 3-several together (up

to 10 on large male stems) on short stout

peduncles, and with a dense yellowish-grey to

brownish woolly covering. Male cones

cylindric, curved, 20-30 cm long, 6-7 cm

diam. ; median scales about 2 • 5-2 • 8 cm long

and 2 cm broad with beak projecting about

5 mm beyond sporangial margin and turned

slightly upwards; bulla face terminating in a

narrow blunt facet under the woolly covering.

Female cones more or less oblong in outline

with rounded apex, 25-30 cm long and 15-17

cm diam. ; median scales 4 • 5-5 cm long ; the

bulla about 4-5 cm broad and 2-5 cm thick

vertically, with short incurved lateral lobes

6-8 mm long; bulla face pale canary-yellow

under the wool, elliptic-rhomboid with more

or less acute lateral ridges; upper and lower

facets broadly rounded, terminal facet flat-

tened, about 1 cm broad and 5 mm wide

vertically. Seeds pale yellow to pale orange-

yellow, 3 cm long and 2 cm diam., angled by

compression. Fig. 4.

Locally frequent on mountain sides near Cathcart

and Queenstown, extending eastwards to the neigh-

bourhood of Kokstad; the more westerly records

are almost certainly incorrect. Map 1.

Cape. — Cathcart: Acocks 10999; Story 2800;

Codd 6356; Dyer 5791; Comins 1841. Queens-

Zamjaceae

11

Fig. 4. — Encephalartos friderici-guilielmi, a, portions of typical leaf, reduced; b, tip ot leaflet, enlarged

(Everitt PRE 29909).

12

Zamiaceae

town: Galpin 1525; 18411; Dyer 5790a; Acocks

18857. Tsolo: Verdoorn & Christian 711; Dyer

4712. Tabankulu: Story 4213.

At different times this species has been confused

with E. cycadifolius Lehm., which, however, is a

smaller plant with more widely spaced leaflets. A

discussion of the problem will be found in the

Journ. S. Afr. Bot. 31 : 116. In Proc. Linn. Soc.

New S. Wales 84 : 107 (1959), Johnson points out

that the epithet pauli-guilielmi is a direct Latin

genitive and the terminal “i” should not be doubled,

which rule applies also in the present case.

4. Encephalartos lanatus Stapf & Burtt

Davy in FI. Transv. 1 : 40, 99, fig. 4D (1926);

Hutch. & Rattr. in F.C. 5, 2 (Suppl.) : 43

(1933) partly, as to specimens from Middel-

burg district (excluding syn. E. laevifolius)',

M. R. Henderson in Journ. S. Afr. Bot. 11:52

(1945) partly, as to specimens from Middel-

burg district. R. A. Dyer in Bothalia 8 : 445

(1965). Type: Transvaal, Middelburg,

Weeber H.6471 (K ex PRE, holo.).

Plants unbranched or sometimes branch-

ed from the base; stems up to about 1 m tall

and only very rarely up to 3 m long and re-

clining, up to about 25 cm diam., with densely

grey woolly bracts. Leaves -6-1 m long, in-

cluding petiole 20-25 cm long, white or grey

woolly when young except under surface of

leaflets, glabescent except base of petiole and

pulvinus; rhachis recurved and often slightly

twisted ; leaflets glaucous when young, entire,

pungent, set in V formation above middle of

rhachis and usually overlapping, reduced in

size to base and apex of rhachis ; median leaf-

lets 10-13 cm long, 6-8 mm broad, finely

10-14-nerved on under surface. Cones 1-4

together on short peduncles, covered by a

dense light cream to greyish tomentum. Male

cones subcylindric, 25-30 cm long, 5-6 cm

diam., narrowed to both ends; median scales

about 2-3 cm long and 2 -5-3 -2 cm broad,

with lateral ridges; bulla face flattened, about

2-5 cm broad and 1-5 cm wide vertically

(with tomentum). Female cones oblong in

outline, 25-35 cm long, 12-15 cm diam.,

slightly narrowed to rounded apex and base;

median scales 3-3 • 5 cm long ; bulla 4 • 3-5 cm

broad, 2 -5-2 -7 cm thick vertically, with in-

curved lateral lobes about 5 mm long, with

acute lateral ridges; bulla with base glabrous,

yellow or greenish-yellow, upper and lower

facets woolly, rounded, terminal facet in-

distinct, about 1-5 cm diam. Seeds yellow,

oblong-orbicular, 2-5-3 cm long, 2-25-2-5

cm broad with scanty pulp.

In sheltered valleys of the upper reaches of the

Olifants River catchment in the Middelburg, Wit-

bank and Bronkhorstspruit districts, sometimes

common locally. Map 1.

Transvaal. — -Middelburg: Weeber H.6471 ; Pole

Evans H.11427; Marloth 11748; Verdoorn 2143;

2144; 2145; 2146; 2467; Dyer & Verdoorn 4411;

Codd & Verdoorn 10106. Bronkhorstspruit: Dyer

photographs; Verdoorn 2259; Van Ryneveld in

PRE 29803.

The key and descriptions reveal the distinguishing

characters between this and the two species E.

humilis Verdoorn and E. laevifolius Stapf & Burtt

Davy, both of which have been confused with it.

Both these latter are restricted to the catchment area

of the Crocodile River while E. lanatus is restricted

to that of the upper reaches of the Olifants River

including the Wilge River and other tributaries.

5. Encephalartos humilis Verdoorn in

Bothalia 6 : 220, pl.3, 241 (1951); R. A.

Dyer in Bothalia 8 : 448 (1965). Type: Trans-

vaal, Nelspruit, Dyer 4806 (PRE, holo.).

Plant dwarf, branching freely from the

parent rootstock forming small clumps;

stems subterranean or very occasionally up

to about 30 cm above ground with a total

length of 35 cm rarely up to 50 cm, 13-18 cm

diam. with short lanate bracts. Leaves 30-50

cm long including petiole 5-15 cm long,

loosely greyish tomentcse with villous leaf-

lets, glabrescent except for base of petiole

and pulvinus; rhachis usually somewhat

recurved and sometimes slightly twisted to-

wards apex; leaflets glaucous when young,

in V disposition above the middle of the

rhachis and sometimes slightly overlapping,

set parallel or only slightly angled to the

rhachis, pungent, entire; median leaflets

9-13 cm long, 4-6 mm broad, finely about

9-nerved on the under surface. Cones single

on a stem, densely greyish tomentose on ex-

posed face. Male cones 15-20 cm long, 4-5

cm broad, narrowed slightly to base and apex

with peduncle up to about 12 cm long; me-

dian scales about 1 • 5 cm long and 2 cm broad,

1-5-1-75 cm thick vertically, with lateral

angles acute; bulla face very little projected,

with terminal facet occupying almost whole

area, hidden by tomentum. Female cones

barrel-shaped, about 27 cm long, 9 cm diam.,

on short peduncle about 3 cm long; median

scale 3 • 5-4 cm long, 4-5 cm broad, 2-2 ■ 5 cm

thick vertically; bulla with lateral ridges ex-

tending into the lateral incurved lobes about

5-10 mm long, rugcse on basal glabrous part;

bulla face nearly flat, upper and lower facets

Zamiaceae

13

rounded, terminal facet about 1 -5 cm broad

and 1 cm wide vertically, sometimes extend-

ing nearly to lower margin. Seeds orange-

yellow, angled by compression, 2-5-3 cm

long, 2-2-5 cm broad.

Occasional on the mountains of the eastern

Transvaal from the Carolina district to Lydenburg

and Nelspruit, usually associated with grass and

often wedged between rocks. Map 1.

Transvaal. — Carolina: Dyer , Verdoorn & Codd

5776; 5777; 5778. Belfast: Smuts 285; Van Biljon

in PRE 29426. Nelspruit: Mogg 17563; Dyer 4805;

4806; Verdoorn 2349 ; 2350; 2351; Liebenberg 335;

Reynolds 3936. Lydenburg: Marais 270; Van de

Ende in PRE 29427; Dyer, Verdoorn & Codd 5787.

The above species concept was included by Hutch.

& Rattr. in F.C. 5, 2 (Suppl.) : 43 (1933) and by

Henderson in Journ. S. Afr. Bot. 11 : 52 (1945)

under E. lanatus Stapf & Burtt Davy. It differs in

its smaller, more consistently branched habit; it is

less woolly and has straighter hairs on the leaflets;

its habitat conditions are different as it occurs in

mountain grassland within the catchment of the

Crocodile River, as opposed to the Olifants River

catchment for E. lanatus.

6. Encephalartos laevifolius Stapf &

Burtt Davy in FI. Transv. 1 : 40 & 99 (1926);

R. A. Dyer in Bothalia 8 : 451 (1965). Type:

Transvaal, Todd s.n. (K. holo. ; in NH 2043,

iso.).

Plants unbranched or branched from

base; stems 1-3 m tall, occasionally up to

3-5 m, 25-30 cm diam., without loose tomen-

tum. Leaves up to about 1 m long, including

petiole about 25 cm long, at first glaucous

and densely pale brown woolly except on

under surface of leaflets, glabrescent except

for tomentose pulvinus; rhachis nearly

straight or recurved and sometimes some-

what twisted above middle ; leaflets spreading

widely in lower half of leaf and in V disposi-

tion above middle, not or only slightly over-

lapping, pungent, entire, reduced in size to

base and apex; median leaflets linear, up to

12 cm long and 5-7 mm broad, rarely up to

1 cm, 10-12-nerved on under surface. Cones

2-4 together on short stout peduncles sub-

tended by linear-lanceolate, densely tomen-

tose bracts, covered by a not very dense whit-

ish tomentum. Male cones subcylindric,

tapering slightly to base and apex, 30-40 cm

long and about 10 cm diam., curved with age;

median scales 3-5-4 cm long, 3 -4-3 -5 cm

broad, shortly stalked, slightly cordate at

base; bulla face projecting about 1 cm,

sharply laterally ridged, with upper facet

more or less humped, lower facet rounded,

terminal facet concave with the lower margin

slightly more prominent than the upper, the

short tomentum decreasing in density and

length towards the almost glabrous base.

Female cones broadly subcylindric,contracted

slightly to the rounded apex and very slightly

to the base, 20-30 cm long, 10-13 cm diam.;

median scales about 4 cm long, 4-4-5 cm

broad and 2-5 cm thick vertically, with in-

curved lateral lobes about 1 • 5 cm long; bulla

face progressively less tomentose to an al-

most glabrous base, sharply laterally ridged,

upper facet rounded but sometimes with 2

slightly prominent ridges, lower facet round-

ed, terminal facet 1-1-5 cm broad and 5-10

mm wide vertically, concave, with the lower

margin slightly more prominent than the

upper. Seed orange-yellow, angled by com-

pression, 2 -5-2 -7 cm long, 2-2-3 cm broad,

with short fleshy apex.

On the Kaapsehoop Mountain of the eastern

Transvaal, in the Nelspruit district and in Swazi-

land; infrequent. Map 1.

Transvaal. — Nelspruit: Thorncroft sub Rogers

28427; Dyer 4803; 4804; Prosser 1479;

Van Noulntys in PRE 10086; Mogg in PRE 29435;

Unkles in PRE 2943 ; Liebenberg 3342.

Swaziland.— Piggs Peak, Compton 32343.

Although the type specimen was recorded as

having been collected ‘near Barberton’ it is probable

that it was actually from the Kaapsehoop Mt., now

included in the Nelspruit district. It is a more

robust species than E. lanatus, with which it has

been confused, and the indumentum of the cones

differs markedly. The distribution areas are ;n

separate river catchments: the Crocodile for E.

laevifolius, where E. humilis and E. paucidentatus also

occur, but under different habitat conditions.

7. Encephalartos inopinus R. A. Dyer in

Bothalia 8 : 169 (1964); Bothalia 8 : 453

(1965). Type: Transvaal, Lydenburg, Dyer,

Codd & Verdoorn 5788 (PRE, holo.).

Plant freely branched from base; stems

up to about 3 m long and often spreading or

reclining at this stage, 17-23 cm diam. (in-

cluding the persistent leaf bases); bracts

lanceolate, about 4-5 cm long, tomentose.

Leaves -8-1 -2 m long including petiole 10-20

cm long; rhachis nearly straight, sometimes

slightly up-curved and twisted towards apex,

glabrous except for woolly back of pulvinus,

flat or ridged on upper surface towards base,

rounded on under surface, 8-13 mm thick;

pulvinus up to 3-5 cm long, 4-5 cm broad,

with dense brown wool on back readily rub-

Zamiaceae

Fig. 5.— Encephalartos inopinus, a, leaf, reduced; b, leaflet {Dyer, Codd & Verdoorn 5788).

Zamiaceae

15

bed off, shrinking in size and wearing down

considerably with age to about 1 • 5-2 cm

long; leaflets glabrous, glaucous on under

surface when young, becoming green with

age, base or foot inserted parallel to axis,

usually pungent, with entire margin or occa-

sionally with 1-2 minute teeth on the lower

margin near and directed towards the apex,

those leaflets above middle of leaf directed

slightly upwards and outwards and sometimes

recurving, not overlapping, reduced in size

near apex of rhachis, those about the middle

spreading more or less at right angles from

rhachis, those below directed slightly down-

wards and slightly falcate, 1-2 cm distant

from each other and reduced in size rather

abruptly to 2-6 prickles on either side of the

rhachis; median leaflets linear-attenuate, 14-

21 cm long, 8-13 mm broad in the lowest ^

and from there gradually tapering to the

slender pungent apex, only slightly narrowed

at base, not very rigid, with 13-21 veins evi-

dent but not raised on the lower surface.

Male cones 2 or more together (immature)

on peduncles about 5 cm long and 1 • 5 cm

thick, 8-10 cm long, 4-4-5 cm diam. ; scales

dense (few showing some dehiscene) about

2 cm long (tips all damaged) 1 -7-2 cm broad,

with acute lateral margin, maximum vertical

thickness 5 mm; microsporangia dense on

lower basal half ; beak projecting about 1 cm.

with upper facet humped in middle, lower

facet nearly flat, glabrous round base of beak,

shortly greyish scurfy-pubescent towards

apex. Female cones not yet recorded. Fig. 5.

On cliffs in hot valleys near the junction of the

Olifants and Steelpoort Rivers in the Lydenburg

district of the Transvaal. Map 1.

Transvaal.— Lydenburg: between Weltevreden

and Krommellenboog Asbestos Mines, Dyer, Codd

& Verdoorn 5788; 5853; 5854; 5855; Louis Naude

in PRE 29805; Van Bit j on in P RE 29753 ; Els photo;

cult. Johannesburg, Van Hoepen in PRE 29663.

This recently described species may prove less

rare than was at first suggested, once the rough

terrain of its natural habitat has been explored more

thoroughly. The downward direction of the lowest

leaflets is more obvious in leaves of young suckers

than those on old trunks.

8. Encephalartos eugene-maraisii Ver-

doorn in Journ. S. Afr. Bot. 11:1 (1945);

M. R. Henderson l.c. 11 ; 39 (1945); R. A.

Dyer in Bothalia 8 : 455 (1965). Type : Trans-

vaal, Waterberg, Marais in Herb. Marloth

13368 (PRE, holo.).

Plant unbranched or more often branch-

ed from base ; stems up to 2 • 5-4 m tall, rarely

more, reclining with age, 30-45 cm diam.

with tomentose bracts. Leaves -7-1 -3 m long

including petiole 16-20 cm long; rhachis

nearly straight with the apical portion slightly

or markedly incurved, glabrous except for

densely tomentose pulvinus mainly hidden by

bracts, glaucous when young; leaflets rigid,

glabrous, glaucous when young, reduced in

size towards apex and base of rhachis; me-

dian leaflets linear-oblong, 15-20 cm long,

1- 3-1 -5 cm broad, cuneate at base, shortly

pungent, entire or occasionally with a single

tooth on lower margin. Cones 1-3 together

on short stout peduncles, dark brown. Male

cones subcylindric, 22-42 cm long 6-8 cm

broad; median scales 2-5-3-25 cm long,

2- 2-5 cm broad, with acute lateral ridges;

bulla face projected into beak 7 mm long,

surface slightly uneven, terminal facet about

3 mm broad and nearly 3 mm wide vertically,

slightly concave. Female cones ovoid to ob-

long-ovoid in outline, 30-50 cm long, 16-20

cm broad; median scales 5-5-5 cm long,

4-5-5 cm broad, 2-5-3 cm thick vertically,

with lateral ridges extending into incurved

lobes about 1 cm long; bulla papillate round

base, rugose in apical half and minutely pube-

scent, upper facet with 1-2 sagittal ridges and

lower facet rounded, terminal facet concave,

1 -5-2 cm broad, slightly less wide vertically.

Seeds amber or sometimes slightly tinged

with red, fleshy, angled by compression,

3- 5-4 cm long, 2-3-3 cm diam.

In isolated mountainous areas of Transvaal, in-

frequent. Map 2.

Transvaal. — Witbank: Dyer, Codd & Verdoorn

5858. Middelburg: Codd & Verdoorn 10111; Mogg

23933. Pietersberg: Codd & Verdoorn 10426.

Waterberg: Marais in Herb. Marloth 13368;

Pole Evans 4457 ; Galpin 13199; 13802; Verdoorn &

Dyer 2211; Dyer 4290; Dyer & Verdoorn 4420;

4421.

The specimens in the Witbank and Middelburg

districts are generally more robust and with

straighter leaves than those from the Waterberg.

There appears no justification for specific distinction

but varietal distinction might be considered war-

ranted by some. Its nearest relatives would appear

to be E. lehmannii Lehm. and E. princeps R. A. Dyer

in the eastern Cape.

9. Encephalartos princeps R. A. Dyer in

Journ. S. Afr. Bot. 31 : 111 (1965); Bothalia

8 : 458 (1965). Type: Cape, Queenstown,

Galpin 8090 (PRE, holo.).

16

Zamiaceae

Map 2. — Distribution map of Encephalartos eugene-

maraisii, E. princeps, E. lehmannii, E. trispinosus

and E. horridus.

E. lehmannii Auct. in part, not of Lehm.; Hutch.

& Rattr. in F.C. 5, 2 (Suppl.) : 36 (1933), as to

specimens from Queenstown, etc.; form “D”

M. R. Henderson in Journ. S. Afr. Bot. 11 : 30(1945).

Plant unbranched or more often branch-

ed from the base with stems up to 3 m tall or

some attaining 4 m when reclining or pro-

cumbent, about 30 cm diam. Leaves 1-1 - 3 m

long including petiole 15-20 cm long, gla-

brous, glaucous when young becoming dark

green with age; rhachis straight for the most

part with recurved and occasionally some-

what twisted apex, and the tomentose pul-

vinus mostly hidden by stem bracts; leaflets

spaced in the lower half of the leaf and re-

duced in size towards the base, often over-

lapping in the upper half of the leaf and in V

disposition; median leaflets up to about 15

cm long and 1 • 3 cm broad, pungent, entire

or very occasionally toothed on lower margin.

Cones 2 or more per crown, dull green, on

short stout peduncles. Male cones subcylin-

dric, narrowing to base and apex, 16-26 cm

tall, 8-10 cm diam.; median scales about 3

cm long, 1 -5-1 -8 cm broad, 8-10 mm thick

vertically ; bulla face projected into a 1 • 2-1 • 5

cm long beak with sharp lateral ridges, ter-

minal facet 5-8 mm broad and 4-6 mm wide

vertically. Female cones more or less barrel-

shaped, about 30 cm long and 20 cm diam.,

with rounded apex; median scales 6-7 cm

long, 4-4-5 cm broad, 3-3-5 cm thick

vertically, with lateral angles extending

into lateral incurved lobes 1-1*5 cm long;

bulla protruding 1-5-2 cm, flattened on

the face, upper and lower facets rounded

or indistinctly ridged, coarsely and irregularly

warty or coarsely papillate and with thin

tomentum of whitish or brownish hairs, ter-

minal facet slightly below centre, rather vari-

able in size, 5 mm-1 • 5 cm broad and • 5-1 cm

wide vertically, rugose, concave with raised

rim. Seeds red, angled by compression, 4—5

cm long, 2 cm diam., fleshy beak about 1-5

cm long.

Limited in distribution in the eastern Cape within

the catchment area of the Kei River and its tribu-

taries, mainly on doleritic outcrops. Map 2.

Cape. — Queenstown: Galpin 8090. Cathcart:

Henderson 1594. King William’s Town: Comins

1661. Komga: Flanagan 1373; Verdoorn &

Christian 708; Dyer 4525; Dyer & Wells 5792;

5793. Tsomo: Sim in K. Butterworth: Verdoorn

& Christian 709; 2421; Dyer 4508; Smith &

Latimer 261; Dyer & Wells 5794; Smit PRE 29907.

? Mquanduli: Smith 5794 (Cult.)

For a better balanced assessment of the relation-

ship between this species concept and that of E.

lehmannii Lehm. and E. trispinosus R. A. Dyer,

further long-term field work is necessary. It is

essential to determine more exact distribution areas,

the degree of variability of characters and especially

cone characters in all groups. Earlier published

views were based on limited material and in-

formation.

10. Encephalartos lehmannii Lehm., Pu-

gill. 6 : 14 (1834); Miq., Monogr. Cycad. 47

(1842); Schuster in Pflanzenr. 4, 1 : 113

(1932), in part, as to typical form; Hutch. &

Rattr. in F.C. 5, 2 (Suppl.) ; 34 (1933), in

part, as to typical form; M. R. Henderson in

Journ. S. Afr. Bot. 11 : 27 (1945), in part, as

to form “A”; R. A. Dyer in Bothalia 8 : 460

(1965). Type: Cape, eastern area, Ecklon &

Zeyher.

Zamia lehmanniana Ecklon & Zeyher in Otto &

Dietr. Allg. Gartenz. 158 (1833); Bot. Zeit. 479

(1833), nomen nudum.

Encephalartos elongatus Lehm. in Tydschr. Nat.

Gesch. 4 : 419, t.8A (1838). Type: Hort. Hamburg.

? E. spinulosus Lehm. in Tydschr. Nat. Gesch. 4,

420 (1837).

Plant unbranched or more frequently

branched from the base with stems up to

about 1 - 5 m tall, rarely up to 2 m, 25—45 cm

diam. Leaves 1-1 - 5 m long including petiole

up to 25 cm long, glabrous, glaucous when

young, dark green with age; rhachis straight,

Zamiaceae

17

or sometimes slightly recurved and rarely with

the apex twisted sideways, with stout tomen-

tose pulvinus with conspicuous red-brown

collar; leaflets spaced in the lower half of the

leaf and reduced in size possibly to one

prickle, closely set in the upper half but not

much overlapping, in V disposition; median

leaflets linear-lanceolate, 12-18 cm long,

1 - 7—1 -9 cm broad, pungent, entire or rarely

with 1 or 2 small teeth on lower margin.

Cones solitary, on short stout peduncles,

blackish-red because of short dense blackish

hairs on the terminal half of the bulla face

or sometimes greyish on the terminal facet.

Male cones subcylindric narrowed to apex

and base, 25-35 cm long, 8-10 cm diam.;

median scales 2-25-2-5 cm broad, 1 cm

thick vertically; bulla face projected into a

beak about 1 • 5 cm long with acute lateral

ridges, terminal facet 5-8 mm broad and

about 5 mm wide vertically. Female cones

ovoid or barrel-shaped, 30-50 cm long,

15-23 cm diam.; median scales about 6-7

cm long and 6 cm broad and 3-5 cm thick

vertically with lateral ridges extending into

the incurved lateral lobes 1-1-5 cm long;

bulla face protruding about 2 cm, almost

smooth, the upper facet with a central ridge,

lower facet rounded, terminal facet 1-5-1 -8

cm broad, 1-1-5 cm wide vertically, slightly

concave with acute upper margin. Seed red,

angled by compression, 4-7-5 cm long, about

2 cm diam., with fleshy beak.

Associated with karroid scrub bush in the Cape

on the Klein-Winterhoekberge and Groot-Bruint-

jieshoogte from near Willowmore to Uitenhage and

Steytlerville to Pearston, Bedford and possibly

northern Albany near Kommadagga. Map 2.

Cape. — Willowmore: Andreae 999; Verdoorn &

Christian 688; 689; 689a; 689b. Steytlerville:

Story 2491. Uitenhage: Dyer 4442; 4443; Dyer &

Wells 5814. Jansenville: Verdoorn & Christian 690;

690a, b, &c; Dyer 4011; Dyer & Wells 5808 ; 5809;

5811; Acocks 11992. Pearston: Jenkins in PRE

29569; Codd 10472. Bedford: Pringle in PRE

29821.

The relationship between E. lehmannii, E. horridus,

E. trispinosus and E. princeps is intimate. Individual

views based on the same facts are unlikely to agree

in detail. There is some discussion in the notes on

E. trispinosus, which is always likely to be a most

controversial concept.

Verdoorn & Christian 689, 689a and 689b are

forms with 1 or 2 prickles of varying sizes on the lower

margin of some of the median and upper leaflets,

jn this they indicate a relationship with E. trispinosus

but the locality is far distant. Other records coming

to hand on going to press indicate that the specimens

cited from Pearston and Bedford may qualify for

varietal distinction from typical E. lehmannii. Female

cone characters may justify this but the available

material is still deplorably meagre. It has been

found that the female cone scales illustrated in

Bothalia l.c. fig. 50, from Groot-Bruintjieshoogte

are smoother than some produced on plants from

the Klein-Winterhoeksberge. The latter show a

similarity to those of E. horridus.

11. Encephalartos trispinosus (Hook.) R.

A. Dyer in Journ. S. Afr. Bot. 31 : 112 (1965);

Bothalia 8 : 463 (1965). Type: Bot. Mag.

t.5371 (1863) represented by specimen in K

(1880).

E. horridus var. trispinosa Hook, in Bot. Mag.

t.5371 (1863). E. horridus sensu Schuster partly in

Pflanzenr. 4, 1 : 116 (1932). E. lehmannii sensu

Hutch. & Rattr. partly in F.C. 5, 2 (Suppl.) ; 34

(1933); M. R. Henderson in Journ. S. Afr. Bot.

11 : 32, Form B1 & C (1945).

Plant branched from the base or some-

times unbranched, with stems up to about

1 m tall and 25-30 cm diam. Leaves usually

glaucous-green when young, becoming green

with age, -75-1-25 m long including petiole

1 5-22 cm long, glabrous ; rhachis stiff usually

recurved and often twisted towards apex,

with dirty white collar round tomentose

pulvinus; leaflets spaced and reduced in size

towards base, occasionally overlapping in

upper half ; median leaflets linear-lanceolate,

or oblong-lanceolate, often somewhat falcate,

10-18 cm long, 1 -5-2-5 cm broad, pungent

with terminal spine up to 6 mm long, usually

with 1-2 pungent lobes 1-3 cm long from

lower margin. Cones solitary, general colour

blue-green or yellowish-green, shortly and

stoutly pedunculate. Male cones subcylindric,

narrowed to both ends, 25-35 cm long,

6 -5-7 -5 cm diam.; median scales about 2-5

cm long, 2 -5-2 -8 cm broad with sharp

lateral angles; bulla face projected into a

beak 7 mm long, terminal facet about 6 mm

broad and 6 mm wide vertically. Female

cones broadly subcylindric, rounded or

narrowed to apex, 40-50 cm long and 16-18

cm diam.; median scales 7-9 cm long, 6-7

cm broad and 3-5-4 cm thick vertically, with

lateral ridges extending into the incurved

lateral lobes 2-3 cm long; bulla face pro-

truding 2-5-3 cm, sparsely hairy with ad-

pressed whitish hairs, upper facet with rugose

ridges converging on the terminal facet and

with irregular pointed papillae round the base,

lower facet rounded, terminal facet smooth

or sparsely grooved, concave, about 1 cm

18

Zamiaceae

broad and 1 cm wide vertically. Seeds reddish

to yellowish-orange, 4—5 cm long and 2 cm

diam., with fleshy beak (there is likely to be

greater variability in cone size than recorded

here based on limited material).

In the eastern Cape in karroid scrub and bush of

the Bushmans, Great Fish River and intervening

valleys. Map 2.

Cape. — Alexandria: Verdoorn & Christian 693;

697; Compton 19697B. Bathurst: Dyer & Wells

5796; 5797; Wells 2904; 2905; Acocks 16152.

Albany: Galpin 3083; Verdoorn & Christian 703;

703a; 704; Dyer 1184; 4842; Dyer & Wells 5801;

Acocks 12769; Smith 5815. Victoria East: Acocks

15979. Peddie: Dyer 4547B; Verdoorn & Christian

705; 706.

Hooker wrote in Botanical Magazine 1863:

“After the most careful examination of our plant

(tab. 5371) I can only come to the conclusion that

it is one of the many varieties of E. horridus ; with

a great tendency to have, on the inferior margin of

the pinnules, two large spinescent laciniae, which,

in conjunction with the terminal spines, justify the

application of the term var. trispinosa” . A herbarium

specimen was preserved at Kew in 1880 but the

original history of the plant was unknown. Schuster

(l.c.) quoted E. horridus var. trispinosa Hook, under

E. horridus (Jacq.) Lehm., while Hutchinson and

Rattray treated the name as a synonym of E.

lehmannii', Henderson also dealt with it under

E. lehmannii, in which he included several forms.

Henderson’s material is reclassified in the present

work under three groups of specific status: E.

lehmannii , E. trispinosus and E. princeps, and a

residue of supposed hybrids of indeterminate origin.

The figure in Bot. Mag. t.537 1 can be matched

very closely with specimens in the lower reaches of

the Bushmans and Fish River Valleys. These speci-

mens appear to be sufficiently distinct, numerous,

widely spread and often isolated from any other

species, to be given specific rank. While there is no

means of confirming that the type specimen was

collected within this environmental range, con-

versely, it would not be possible to deny the pro-

bability. There appears to be no earlier name which

could, with equal confidence, be applied. The reason

for caution in elevating the concept of trispinosus to

specific rank is that a number of specimens within

the same distribution area do not conform to a

species pattern and give the impression that natural

hybridization has taken place. E. horridus var.

trispinosa Hook, first had to be viewed from this

angle.

The first botanist to generalize on the relation-

ships of specimens in the eastern Cape Province was

Karl Zeyher whose notes were read in 1852 before a

scientific gathering and published in the Phytologist.

After mentioning that any person with “tact”

should have no difficulty in detecting the “play of

nature” he went on to say that he intended to

examine all the populations once more in their

natural state. The outcome is not recorded and

Zeyher’s warning seems to have gone unheeded.

Plants of supposed hybrid origin occur two miles

up the Bushmans River and in the lower reaches of

the Fish River Valley. These appear to combine the

characters of E. trispinosus and E. altensteinii both

of which are present in the same areas. E. arenarius

R. A. Dyer, the typical form of which occurs a few

miles west of Bushmans River in a secluded valley,

may also be involved in hybridization, and E.

latifrons cannot be ignored. (There is a strong

suspicion that E. latifrons has been involved in

hybridization with E. altensteinii at least).

The distinction between E. trispinosus and E.

horridus is based largely on the structure of the

female cone scales, for which view relatively few

cones have been available. The distinction between

E. trispinosus and E. lehmannii is based on female

cone scales and leaf characters, cones again being

in short supply. Nevertheless, on the evidence

available it seems that neither typical E. horridus

nor E. lehmannii occurs in the area of distribution

of E. trispinosus. E. horridus is apparently restricted

in its distribution range to the districts of Uitenhage

and Port Elizabeth, while E. lehmannii is in Uiten-

hage, Willowmore and Jansenville, mainly on the

lower slopes of the Klein-Winterhoekberge, with a

form in the Pearston, Somerset East and Bedford

districts on the Groot-Bruintjieshoogte. It is possible

that it occurs also in northern Albany district near

Kommadagga.

A study of these three species has led to the

suggestion that both E. horridus and E. lehmannii

evolved from E. trispinosus stock. The stem has

been reduced and the lobing of the leaflets accen-

tuated in E. horridus, whereas in E. lehmannii the

stems are more robust and the leaflets are mainly

entire. An occasional part-reversion within the

normal populations of these two species in the

direction of E. trispinosus indicates this possibility —

a stem up to 4 ft in E. horridus ( Story 2338) and

specimens of E. lehmannii with teeth on the lower

margin of some leaflets ( Verdoorn & Christian 689,

689a, 689b). It is believed that these variations have

been viewed on a rational basis in the present

treatment in not giving them distinctive rank. Were

cones available from all specimens it is felt that

they would be sufficiently distinctive within the

groups to maintain the present basic classification.

Of the herbarium material examined from known

localities, the following show a strong affinity with

E. trispinosus but leave some doubt about their

complete identity:

Bathurst: Southwell, Acocks 16152. Peddie:

Dyer 4547A & C. (into this category Henderson,

l.c., Forms B, B2 and C also fall.)

The following appear to be of hybrid origin

involving E. altensteinii as the other parent :

Alexandria: Bushmans River, Verdoorn &

Christian 693; 695; 696; 698; 699; Archibald

4342 A; 5590; Compton 19697A. Bathurst: Dyer

& Wells 5800; Wells 2901 (nearer E. altensteinii)-.

Wells 2902 (nearer E. trispinosus).

The following appear to be of hybrid origin

involving E. arenarius as a parent:

Bathurst: Southwell, Acocks 16153; Dyer 4835.

Zamiaceae

19

The following single specimen found with E.

altensteinii (but out of the proved range of E.

trispinosus , and possibly since drowned in Murray

Reservoir in the King William’s Town district) sets

a problem which cannot be followed further at this

stage.

King William’s Town: Smith 6064 (= Verdoorn

& Christian 707 and Dyer 4847 in Hort., East

London).

As we go to press a leaf, not unlike that of Smith

6064, but whose leaflets have no prickles, has been

brought to the Botanical Research Institute, Pre-

toria, by Mr. J. A. Smit (PRE 29906) from about 9

miles from the Keiskamma River Mouth in the

Peddie district. This is between the Bathurst and

King William’s Town districts. In other circum-

stances it might well have been identified as E.

lehmannii, but it is suspected that the female cones,

when recorded, will show a closer affinity with those

of E. trispinosus.

12. Encephalartos horridus ( Jacq .) Lehm.,

Pugill. 6 : 14(1834); Schuster in Pflanzenr. 4,

1 : 116 (1932), in part, excluding varieties;

Hutch. & Rattr. in F.C. 5, 2 (Suppl.) : 32

(1933); M. R. Henderson in Journ. S. Afr.

Bot. 1 1 : 20 (1945); R. A. Dyer in Bothalia

8 : 467 (1965). Type: Cape, without locality,

the illustration in Jacq. Fragm. 27, t.28 (1801)

from a plant cultivated in Vienna, leg. Scholl.

Zamia horrida Jacq. Frag. 27 : t.28 (1801),

probably excluding t.27. ? Z. gleina Miq. in Linnaea

17 : 729 (1843). ? Z. aurea Hort. ex Miq. in Tyd.

Wet. 1 : 297 (1848).

Plant unbranched or more usually

branched from the base with stems subterra-

nean or partly exposed, up to about 30 cm

tall, exceptionally up to 1 m tall, 20-30 cm

diam., the crown with a small quantity of

loose wool. Leaves several to many in a whorl,

rigid, glaucous, glabrous, -5-1 m long in-

cluding petiole up to about 13 cm long;

rhachis much recurved in the apical third,

with brown woolly pulvinus; leaflets rather

widely spaced in lower half of leaf, closer in

the upper half, reduced in size and entire to-

wards base; median leaflets ovate to lanceo-

late in general outline, up to 10 cm long and

2-5 cm broad with pungent apex and 1-3

pungent lobes up to 4 cm prominent on lower

margin and twisted out of the plane of the

leaflet; nervation obscure. Cones solitary on

short stout peduncles, with blackish or

reddish-brown adpressed scurfy hairs over

green surface. Male cones subcylindric nar-

rowing gradually to base and apex, 20-40 cm

long, 6-12 cm diam.; median scales broadly

ovate, 3-4 cm long and 3-3 • 5 cm broad, with

acute lateral ridges; bulla face projecting

8-10 mm beyond sporangial surface, smooth

or slightly corrugated, upper facet humped

and sometimes with obscure ridges, terminal

facet concave, smooth, about 8 mm broad and

6 mm wide vertically. Female cones ovate-

oblong to oblong-elliptic in outline, 20-40 cm

long, 1 5-20 cm diam., rarely smaller; median

scales 4-5-6 cm long, 4-5 cm broad and

3-3-5 cm thick vertically, with lateral ridges

extending into the incurved lateral lobes

about 1 cm long; bulla face protruding 1 -5-2

cm, rugose round base, upper facet with 1-2

or more ridges converging to the terminal

facet otherwise fairly smooth, lower facet

slightly protuberant, somewhat rugose, termi-

nal facet more or less quadrangular, 1 ■ 5-2 cm

broad, concave with slightly uneven surface

and a narrow raised rim. Seed pale red or

carmine, 3-3-5 cm long and about 2-5 cm

diam., angled by compression, with fleshy

beak.

Local in the Port Elizabeth and Uitenhage dis-

tricts of the eastern Cape. Map 2.

Cape.— Port Elizabeth: Dyer 5060; Rodin 4833.

Uitenhage: Verdoorn & Christian 680; 681; Dyer

4439; Smith 3; Story 2338 (form with stem up to

4 ft long); Dyer & Wells 5818.

A small group of plants from infertile sourveld

on quartzite hills west of Port Elizabeth are dwarf

by comparison with those from the more fertile

but drier karroid scrub bush in the Uitenhage

district. In the latter area specimens may, on rare

occasions, reach a height of 4 ft whereas the stems

of others may be completely subterranean. Jacquin

described the type as with a stem 2 ft long but it is

not known how much of it was originally below

ground. The Fig. 27 of the cone in Jacquin’s

Fragmenta is not good as regards detail and may

have originated from E. longifolius. These two

species have been found growing together near

Uitenhage, but there was no suggestion of inter-

specific hybridization at the time.

13. Encephalartos arenarius R. A. Dyer in

Journ. S. Afr. Bot. 22 : 1 (1956); Bothalia

8 : 470 (1965). Type: Cape, Alexandria Dis-

trict, Dyer 5441 (PRE, holo.).

Plant unbranched or more usually

branched from the base; stem up to about 1

m tall and at this length often somewhat re-

clinate, 20-30 cm diam. Leaves 1-1 -5 m long

including petiole 15-20 cm long; rhachis

spreading, recurved in upper half, glabrous

except for brown tomentose pulvinus; leaflets

somewhat glaucous, light green when young

gradually darker green with age, overlapping

20

Zamiaceae

in the apical half and in V disposition, more

spaced in the lower half and reduced in size

occasionally to one prickle; median leaflets

oblong-lanceolate, flat or slightly twisted and

slightly curved upwards, 12-16 cm long,

2 • 5-4 cm broad with 1-1 • 5 cm broad attach-

ment, closely but not prominently veined on

lower surface, pungent with upper margin

entire or occasionally a single tooth and lower

margin usually with 3 rarely 4 flat or slightly

twisted pungent lobes. Cones single on short

stout peduncles. Male cones (incomplete);

median scales 3-3-5 cm long, 2-25-2-5 cm

broad ; bulla face projected into a beak about

1 cm long. Female cones mainly barrel-

shaped with rounded base and apex, 35-45 cm

long and 20 cm diam., somewhat glaucous;

median scales about 6 • 5-7 cm long, 5 cm broad

and 4 cm thick vertically, with lateral ridges

extending into incurved lateral lobes up to 2

cm long; bulla face projecting about 2 cm,

upper and lower facets with or without ridges

and surface slightly and irregularly undulate

or minutely papillate; terminal facet 10-12

mm broad and about the same in width verti-

cally, concave. Seeds coral-red, up to about

5 cm long, 2-2-5 cm broad with fleshy beak.

Local in the Alexandria District of the Cape,

associated with coastal sand-dune scrub bush.

Map 3.

Map 3. — Distribution map of E. arenarius and E.

latifrons.

Cape. — Alexandria: Dyer 5439; 5441; 5443;

Gledhill 5420; Verdoorn 2422; Guillarmod 4302.

Notes under E. trispinosus and E. altensteinii state

that this species is possibly involved in hybridization

with those species but neither occurs in the type

locality of E. arenarius in the Caba Valley. It is

nearest related to E. latifrons.

14. Encephalartos latifrons Lehm. in

Tydschr. Nat. Gesch. 4 : 424, t.9 A & B

(1837-38); Hutch. & Rattr. in F.C. 5, 2

(Suppl.): 36 (1933); M. R. Henderson in

Journ. S. Afr. Bot. 11 : 22 (1945); R. A. Dyer

in Bothalia 8 : 471 (1965). Type: the illus-

tration in Tydschr. Nat. Gesch. 4 : t.9

(1837-38).

E. horridus var. latifrons Miq., Monogr. Cycad.

59 (1842); Schuster in Pflanzenr. 4, 1 : 177 (1932).

Plant unbranched or more usually

branched from the base with stems up to

about 2 • 5-3 m tall, rarely taller. Leaves finely

pubescent throughout when young, gla-

brescent with age, 1-1-5 m long including

petiole 10-20 cm long; rhachis rigid, recurved

or curled in the upper half, thickly tomentose

round base and on pulvinus with whitish

collar; leaflets rigid, dark green, overlapping

in the upper half of leaf and in V disposition,

more spreading in lower half of leaf and

gradually reduced in size, but only the lowest

sometimes prickle-like; median leaflets ovate-

oblong, 10-15 cm long, 4-6 cm broad with

attachment 1 • 5-2 cm broad, pungent,

with upper margin entire, rarely toothed,

lower margin with 2-4 deep triangular pun-

gent lobes sometimes somewhat twisted.

Cones 1-3 together on short stout peduncles,

general colour dark green or dark blue-green.

Male cones subcylindric narrowed to both

ends, 30-50 cm long, 8-17 cm diam. ; median

scales 6-7 cm long, broadly obovate-oblong

in the sporangial portion, 4-4-5 cm long,

3-3-5 cm broad, with acute lateral ridges;

bulla face projecting into a slightly decurved

beak about 2 cm long, sparsely covered with

adpressed red scaly hairs, upper and lower

facets variably ribbed, terminal facet slightly

concave, about 1 cm broad and slightly nar-

rower vertically. Female cones barrel-shaped,

up to about 60 cm long and 25 cm diam.;

median scales about 8 • 5 cm long and 5-5 cm

broad, with lateral ridges extending into in-

curved lateral lobes about 1 cm long; bulla

face protruding 2-2-5 cm, the surface with

coarse pointed or blunt papillae and pale

brownish scaly hairs mixed with scales, upper

facet ridged or crested; lower facet rounded;

terminal facet irregularly shaped, 1-1-3 cm

diam., slightly narrower vertically, concave or

excavate with prominent margin. Seeds red,

angled by compression, about 5 cm long and

2-2-5 cm diam., with fleshv beak Fig. 6.

In association with scrub bush on rocky outcrops

in the Bathurst and Albany Districts of the eastern

Cape; rare. Records from outside this area require

verification. Map 3.

Zamiaceae

21

Fig. 6. — Encephalartos latifrons, a, middle portion of leaf, much reduced; b, leaflet showing venation; both

reduced (taken from F.C. 5, 2 (Suppl.) 1933.)

22

Zamiaceae

Cape. — Bathurst: Galpin 8439; Verdoorn &

Christian 702; Dyer 4556; 4561. Albany: Story

2815; 4540.

This is one of, if not the slowest growing arbore-

scent species in South Africa. It seems that intervals

of at least 2 or more years elapse between the pro-

duction of new whorls of leaves.

A specimen in the garden on the farm Wylmington

between Fish River and Shaw Park ( Dyer 4559),

strongly suggests hybridization between E. latifrons

and E. altensteinii, as does another in the garden of

the Albany Museum, Grahamstown. There seems no

good reason for the confusion which has arisen at

times between this species and E. horridus. The

MacOwan specimen cited in F.C. from the Uitenhage

District indicates an error in labelling.

15. Encephalartos longifolius ( Jacq .)

Lehm., Pugill. 6 : 14 (1834); Schuster in

Pflanzenr. 4, 1 ; 111 (1932); Hutch. & Rattr.

in F.C. 5, 2 (Suppl.): 38 (1933); M. R. Hen-

derson in Journ. S. Afr. Bot. 11 : 40 (1945);

R. A. Dyer in Bothalia 8 : 474 (1965). Type:

Cape, without locality, the illustration in

Jacq., Fragm. 28, t.29 (1801) from a plant

cultivated in Vienna, leg. Scholl.

Zamia longifolia Jacq., Fragm. 1 : 28, t.29 (1801).

Z. lanuginosa Jacq., Fragm. 1 : t.30 & 31 ( 1 801).

Syntypes: Cape, Jacquin’s illustrations. ? Z.

elegans Yates in Proc. Linn. Soc. 2:18 (1849).

Cycas caffra Thunb., in Nor. Act. Soc. Sc.

Uppsal. 2 : 283 (1775) in part, as to part of descrip-

tion and part of Thunberg’s material at Uppsala,

excluding t.5.

Encephalartos caffer sensu Hook, in Bot. Mag.

82 : t.4903. E. lanuginosus (Jacq.) Lehm., Pugill.

6 : 14 (1834).

Plants unbranched or branched from the

base with stems up to about 3 m tall, occasion-

ally up to even 4 • 5 m, 30-45 cm diam. Leaves

1—1 * 75 m long including petiole 15-35 cm

long; rhachis spreading erect, recurved to-

wards apex or arcuate, finely pubescent, gla-

brescent with age except for woolly pulvinus;

leaflets dark green or dark green with bluish

lustre, finely pubescent, glabrescent with age,

rigid in texture, overlapping in the upper half

of the leaf and disposed in V formation, re-

duced in size towards base, sometimes to one

or two prickles; median leaflets mainly

oblong-lanceolate up to 20 cm long and 4 cm

broad, pungent or blunt, entire for the most

part or with the lower margin (mainly of up-

per leaflets) with 1-3 short pungent or blunt

teeth. Cones 1 or 2 on short stout peduncles,

greenish-brown with reddish adpressed hairs.

Male cones subcylindric, narrowed somewhat

to the ends, 40-60 cm long and 14-20 cm

broad ; median scales up to about 4 cm broad ;

bulla projecting into a 2 cm long beak with

lateral ridges, the upper and lower facets

humped towards apex, the terminal facet

8-9 mm broad and 3-5 mm wide vertically.

Female cones ovid, or barrel-shaped, about

60 cm long and 40 cm diam.; median scales

7 • 5-8 • 5 cm long, 5 • 5-6 cm broad, 4-4 • 5 cm

thick vertically with lateral ridges extending

into incurved lateral lobes about 2-5 cm long;

bulla projecting 2-2 • 5 cm, finely rugose round

the base, coarsely rugose in the apical half,

upper facet with central ridge or hump, lower

facet more or less rounded, terminal facet

about 1 • 5-2 cm broad and the same vertically.

Seeds red, about 5 cm long, 2-5 cm diam.,

with fleshy beak.

At intervals on exposed mountain slopes from

Uniondale to Albany District, sometimes locally

frequent. Map 4.

Map 4. — Distribution map of E. longifolius, E.

altensteinii, E. rvoodii, E. natalensis, E.

lebomboensis, E. transvenosus, E. paucidentatus

and E. ferox.

Cape. — Uniondale: Joubertina, Dyer & Wells

5805; 5807. Humansdorp: Dyer 4447; Story 2457;

Dver & Wells 5804. Uitenhage: Verdoorn &

Christian 676; 679; 682; 682a; 683; 684; 685;

686; 691; 691a; Dyer 4449; 4451. Somerset East :

Verdoorn & Christian 692; 692a. Albany: Dyer

3328; 5449; Wells 2893.

The cones of this species are among the largest

in the genus and may weigh up to about 70 lb.

In some relatively young specimens, particularly in

the most westerly area of distribution, the leaves

Zamiaceae

23

are decidedly glaucous, giving the impression that

two distinct species are present, but field observa-

tions near Joubertina indicate that it is merely

infraspecific variability. The nearest situated

normally glaucous-leaved species to that site is

E. lehmannii which is separated by the Langeberge.

These two species occur fairly near together at the

northern entrance to Paardepoort but evidence of

hybridization has not been found there. E. longi-

folius has recently been found in association with

E. horridus on a hill near Uitenhage but no hybri-

dization between the two species was observed there

either. Its nearest related species is E. altensteinii.

16. Encephalartos altensteinii Lehm.,

Pugill. 6 : 11, t.4 & 5 (1834); Hook.f. in Bot.

Mag. 117 : t.7162, 7163 (1891), excluding

concept of E. natalensis Dyer & Verdoorn;

Marloth, FI. S. Afr. Fig. 62, 64, t.15, 16a

(1913); Schuster in Pflanzenr. 4, 1 : 112

(1932); Mutch. & Rattr. in F. C. 5, 2 (Suppl.):

39 (1933), in part; M. R. Henderson in Journ.

S. Afr. Bot. 11 : 41 (1945), all excluding con-

cept of E. natalensis', R. A. Dyer in Bothalia

8 : 477 (1965). Type: Cape t.4 in Pugill l.c.

leg. Ecklon & Zeyher.

E. marumii de Vriese in Tydschr. Nat. Gesch.

5 : 188 (1838). Type: Hort. Amsterdam.

Plants branched or unbranched with

stems up to about 4 m tall, rarely up to 7 m

and then usually reclining, 25-35 cm diam.

with small amount of wool at apex soon dis-

appearing. Leaves 1-2 m long including

petiole 10-30 cm long, tomentose on emer-

gence except cn upper surface of leaflets soon

glabrescent; rhachis stiff and nearly straight

when in exposed habitat, curved in shade,

glabrous except for pulvinus; leaflets vari-

ably rigid, reduced in size to base but not to

a series of prickles, inconspicuously nerved;

median leaflets oblong-lanceolate to lanceo-

late up to about 1 5 cm long and 2 • 5 cm broad,

with 1 or 2 or, less frequently, 3-5 teeth on

upper and lower margin, sometimes on one

margin and not the other, rarely entire,

mucronate. Cones 2-5 together, rarely only

1, on short stout peduncles, yellowish. Male

cones subcylindric, obtuse or narrowed to

apex, up to about 50 cm long, 1 2 cm in diam. ;

median scales up to 5 cm long and 3 cm broad

with acute lateral ridges; bulla projected

into a decurved beak about 1-5 cm long,

upper facet with central ridge, lower facet

rounded, terminal facet flat or slightly con-

cave, about 1 cm broad and 5-10 mm wide

vertically with sparse mixed reddish and white

scaly hairs. Female cones broadly subcylin-

dric, obtuse or narrowed to apex, 40-55 cm

long, 20-28 cm diam.; median scales 7-8 cm

long, 8-9 cm broad, 3 • 5-4 cm thick vertically,

with lateral ridges extending into incurved

lateral lacerate lobes about 3 cm long and

almost reaching the cone axis; the bulla face

deeply wrinkled and 2-3 cm prominent. Seeds

scarlet, angled by compression, 3-5-4 cm

long, 2-2-25 cm diam., with fleshy beak.

Coastal districts of the eastern Cape from Bush-

mans River to Transkei, associated with low forest

but sometimes on exposed cliffs. Map 4.

Cape. — Alexandria: Copeman in PRE 29556;

Verdoorn & Christian 699. Bathurst: Verdoorn &

Christian 700; Dyer 4555; 4557; 4558. King

William’s Town: Smith 5771; 6873; Dyer 4843;

4844; Acocks 11876. East London: Galpin 7104;

Dyer 4428; 4846. Komga: Flanagan 1372. Ken-

tani: Pegler 116. Elliottdale: Story 4123. Lusi-

kisiki: Crundall in PRE 29557. This last specimen

of an incomplete leaf has leaflets much smaller than

average but possibly represents an outlying form.

A specimen received from the Botanical Garden

of Hamburg in 1952 closely matches the type figure.

The parent plant from which it was taken may

possibly have been there in Lehmann’s time.

Specimens within this distribution area, which

have prickles on the petiole, suggest hybrid influence

with E. villosus Lem. There is strong evidence for

this supposition near East London. Such specimens

could be mistaken for forms of E. natalensis, the

concept of which has been associated with E.

altensteinii by several writers.

There is overwhelming circumstantial evidence to

suggest that natural hybridization is not uncommon

between E. altensteinii and E. trispinosus when they

occur together in the wild state (see notes under the

latter).

A specimen in cultivation in the Bathurst District,

Dyer 4559 and less convincingly another in the

grounds of the Albany Museum, Grahamstown,

Wells 2907, suggest hybridization between this and

E. latifrons.

17. Encephalartos woodii Sander i n Gard.

Chron. 1908: 257 (1908); Praia in Kew Bull.

1914 : 250 (1914) with habit fig., and 1916 :

181 (1916) in text; Schuster in Pflanzenr. 4,

1 : 120 (1932); Hutch. & Rattr. in F.C. 5,

2 (Suppl.): 40 (1933: M. R. Henderson in

Journ. S. Afr. Bot. 1 1 : 47 (1945); R. A. Dyer

in Bothalia 8 : 482 (1965). Type: Natal,

Zululand, Medley Wood , photograph in

Sander, l.c.

E. altensteinii var. bispinosa J. M. Wood in Ann.

Rep. Durban Bot. Gard. 1907: 8, with fig. Type:

Wood, cult. NH.

Plant (only male known) branching from

the base (sometimes branched above in cul-

2-F.S.A.

24

Zamiaceae

tivation) ; stem up to 6 m tall, 50-90 cm diam.

at base, narrowing to about 50-60 cm above,

with densely leafy crown and woolly bracts.

Leaves 1-8-2 -5 m long including petiole 10-

20 cm long; rhachis gracefully spreading-

recurved or arched, densely pale brown wool-

ly at first becoming glabrous with age except

for large pulvinus about 10 cm broad; leaf-

lets leathery, dark green, gradually reduced

to prickles at the base, those from juvenile

suckers variably 2-5-toothed in the lower

half on one or both margins, those on old

stems often with entire margins; median

leaflets more or less ovate-lanceolate, up to

about 20 cm long and 5 cm broad, the apex

often slightly hooked and blunt. Male cone

1 -several, bright orange-yellow, on short

stout peduncles, subcylindric, 40-90 cm long,

rarely up to 120 cm, 15-20 cm diam., median

scales up to about 7 cm long and 3-3 • 5 cm

broad; bulla with the beak projecting 2-5-

3 • 5 cm beyond sporangial surface, with acute

lateral ridges, the upper facet with a central

ridge or hump, the lower facet rounded or

slightly ridged, the terminal facet slightly

concave, about 1 cm broad and about the

same vertically, often with raised margin.

Recorded from one locality only near Ngoye in

Zululand and possibly now extinct in the wild;

recorded in the wild by Chamberlain in 1912 and

lastly by a Forest Officer in 1916: see R. A. Dyer

in Bothalia 8 ; 485 (1965). Several plants are in

cultivation locally and overseas. Map 4.

Natal. — Mtunzini: Wylie in NH 16044; 16046;

(Cult. Durban Bot. Card.) Medley Wood ex Forbes

in PRE 29381; Dyer & Verdoorn 2361 ; 2362; 2363.

Our knowledge of E. woodii is based on suckers

of different ages from a single male specimen, the

rootstock of which appears to have perished,

leaving only its transplanted progeny in cultivation.

The female form is unknown. One may speculate

that the single male plant was a relic of a species

now extinct or it could possibly have originated as

a mutation within the concept described as E.

natalensis Dyer & Verdoorn. This latter suggestion

is contained indirectly in Medley Wood’s original

observation that it was possibly a variety of E.

altensteinii Lehm., the name under which E. natal-

ensis was then known. The broadening of the stem

base with age into a form of buttress and the long,

gracefully curving leaves with a densely woolly

young rhachis are among the main distinguishing

characters of the old plants in cultivation.

18. Encephalartos natalensis Dyer &

Verdoorn in Bothalia 6 : 205 (1951); R. A.

Dyer in Bothalia 8 : 486 (1965). Type; Natal,

Dyer 4475 (PRE, holo.).

E. altensteinii Auct., all in part, not of Lehm.;

including Gard. Chron. 6 : 392-97 fig. 80-83 (1876);

ser. 3, 2 : 281 (1887); Hutch. & Rattr. in F.C. 5,

2 (Suppl.) : 39 (1933); M. R. Henderson in Journ.

S. Afr. Bot. 11 : 41 (1945).

Plants branched from the base or occa-

sionally unbranched, with stems 3^4 m tall

and occasionally up to 6 • 5 m, 25-40 cm diam.,

sometimes showing small amount of wool

round apex. Leaves 1 • 3-3 • 2 m long including

petiole 10—26 cm long; rhachis erect spread-

ing, sometimes slightly recurved and twisted

towards apex, tomentose at first, glabrescent

with age except for densely woolly pulvinus;

leaflets mainly spreading in the lower half of

leaf and gradually in narrower V disposition

towards apex, reduced to several prickles

towards base of rhachis, dark green; median

leaflets lanceolate to ovate-lanceolate, lb-

23 cm long, 2 -5-4 -5 cm broad, pungent, en-

tire or with 1-5 sharp prickles on one or both

margins, more often on the lower leaflets

and sometimes almost lobate on young

plants. Cones 2-3 together on stout peduncles,

dark yellowish-green with thin foxy tomen-

tum. Male cones up to about 45 cm long,

9-10 cm diam., median scales about 4 cm

long, 2-5-3 cm broad with sporangial surface

obovate-oblong; bulla extended into a beak

about 1 • 5 cm long with acute lateral ridges,

the upper and lower facets fairly smooth,

terminal facet about 8 mm broad, 7 mm wide

vertically and with raised lower margin.

Female cones oblong or oblong-ovate in out-

line up to 50 cm long and 25 cm diam.; me-

dian scales 7-8 cm long, 5 cm broad and 4 cm

thick vertically with lateral ridges extending

into the incurved lateral toothed lobes 2-3

cm long; bulla face about 2 cm prominent,

finely rugose round base with upper and lower

facets rounded or humped and with promi-

nent blunt rugosites, terminal facet about 2

cm broad and 1 • 5 cm wide vertically, slightly

concave. Seed scarlet, 5 cm long and 2 cm

diam., with fleshy beak.

On cliffs in midland areas from the eastern Cape

border to northern Natal; associated with but

rarely within forest; occasional. Map 4.

Cape. — Tabankulu, Story 4219.

Natal. — Port Shepstone: Oribi Gorge, Dyer &

Dohse 5412; 5418; 5420; 5421; 5422; Beard

724. Pinetown: Edwards 2865. Camperdown:

Dyer 4475; 4476; 4477; Dyer & Verdoorn 2367;

2368; 2369; 2371. Lions River: Bayer S7; Dyer

& Codd 4785. Krantzkop: Verdoorn & Christian

714; 714a. Vryheid: Acocks 11689; Dyer 4696;

Zamiaceae

25

4697; Ward 3398. Ngotshe: Codd 9568; Gerstner

5144.

There is no difficulty in distinguishing typical

E. natalensis from E. altensteinii by the pubescence

and series of prickles towards the base of the leaf

rhachis of the former and their absence from the

latter; the leaflets of the typical forms differ also

but it is not known if the characters merge in the

eastern Cape-southern Natal area. Assumed

hybrids between E. altensteinii and E. villosus near

East London show a strong likeness to some forms

of E. natalensis in Natal. E. villosus has not been

recorded in the presence of E. natalensis in the wild.

1 9. Encephalartos lebomboensis Verdoorn

in Flow. PI. Afr. 27: PI. 1078, 1079 (1949);

R. A. Dyer in Bothalia 8 : 491 (1965). Syn-

types: Swaziland, Keith in PRE 28362 a & b

(male and female respectively).

Plant branched or unbranched from the

base with stems up to about 4 m tall and 30

cm diam., somewhat woolly on crown, with

linear-lanceolate bracts about 4 cm long, 1

cm broad and woolly on back. Leaves in a

dense crown, 1-2 m long, rarely longer,

including petiole 3-10 cm long, pubescent

when young, glabrescent; rhachis almost

straight except towards tip which may be

recurved or twisted, brown woolly towards

base when young, glabrescent except pul-

vinus; leaflets mainly spreading widely,

crowded above middle of rhachis but not

much overlapping, wider spaced towards

the base of rhachis and gradually reduced to

several prickles; median leaflets narrowly

lanceolate or sometimes oblong-lanceolate,

12-17 cm long, 1 -2-2-2 cm broad, with 1-4

prickles on both margins, more regularly on

the lower margin, rarely without prickles.

Cones 1-3 together on short stout peduncles,

apricot-yellow or dull salmon-pink. Male

cones up to about 45 cm long and 13 cm in

diam., narrowed gradually to apex and base;

median scales about 3 • 5 cm long, 4 cm broad

and 1 • 5 cm thick vertically, with sharp lateral

irregularly laciniate ridges; bulla projected

into a beak 1-1-5 cm long, terminal facet

about 1 cm broad and slightly narrower

vertically, with acute lower margin and in-

distinct upper one. Female cones cylindric-

ovoid, up to about 45 cm long and 22 cm

diam.; median scales about 6 cm long, 4-8

cm broad and 3 • 5 thick vertically, with lateral

ridges extending into the incurved lateral

irregularly laciniate winged-lobes about 1-8

cm long; bulla face protruding about 1 cm,

with greyish or foxy pubescence, upper and

lower facets more or less rounded and rarely

ridged, the terminal facet central, slightly

concave, 10-20 mm broad and 5-10 mm wide

vertically, tomentose. Seeds scarlet, 4 cm

long, 1-8-2 -2 cm diam., with fleshy beak.

On cliffs and in gorges of the Lebombo Mountain

range in Swaziland and extending into the south-

eastern Transvaal, northern Natal and Mozambique.

Map 4.

Natal. — Paulpietersburg: Schoeman 1; Ver-

doorn 23lO\ 2311; Dyer & Codd 5032; 5033; 5034.

Ngotshe: Ward 3919. Ubombo: Codd & Verdoorn

10282. Ingwavuma: Verdoorn & Christian 717;

718a; 719; 721; West 2118; Codd & Verdoorn

10300; 10301; Ward 4\Q6\ 4107.

Transvaal. — Piet Retief: Du Plessis 114a; 204;

Dyer & Verdoorn 5837.

Swaziland. — Stegi, Keith in PRE 28352a, b;

Verdoorn & Christian 723; 724; Dyer 4193', 4795;

Compton 29292.

E. lebomboensis meets E. villosus inland from

Pongola Poort in the Transvaal (and probably

elsewhere in the valley) and plants of suspected

hybrid origin {Dyer & Verdoorn 5838) were observed

in close proximity to the two species. At Pongola

Poort on the Transvaal side of the Jozini Dam wall,

E. ngoyanus occurs with E. lebomboensis but evidence

of hybridization between these species is lacking.

The closest relationship of E. lebomboensis in

South Africa is with E. natalensis. Although prickles

on the rhachis, nearly to the base, are a feature of

E. lebomboensis and E. natalensis, occasional speci-

mens in the Paulpietersburg district may be almost

devoid of them as in E. altensteinii. This observation

presents a problem for further investigation. The

female cone scales seem tolerably uniform and do

not show the prominent wrinkled face of E. natalensis

or E. altensteinii.

20. Encephalartos transvenosus Stapf &

Bwlt Davy, in Burtt Davy, FI. Transv. 1: 40,

99, Fig. 4 B (1926); M. R. Flenderson in

Journ. S. Afr. Bot. 11 : 45 (1945); Miller in

Journ. Bot. Soc. S. Afr. 30 : 14 (1944); R. A.

Dyer in Bothalia 8 : 494 (1965). Type:

Transvaal, Letaba, Burtt Davy H. 3086

(PRE, holo.).

E. altensteinii sensu Hutch. & Rattr. in F.C. 5, 2

(Suppl.) : 39 (1933), partly.

Plant generally unbranched, occasionally

branched after injury to crown, maximum

height about 13 m (35-40 ft, many between

15-25 ft), 40-45 cm diam., apex of stem with

some brown wool. Leaves 1-2-5 m long with

fairly short petiole, densely tomentose at first,

glabrescent, except for pulvinus; rhachis

mainly straight, recurving towards apex;

26

Zamiaceae

leaflets spreading-recurved more or less at

right angles to rhachis, upper ones often

somewhat overlapping, reduced in size to-

wards apex and to several prickles at base;

median leaflets lanceolate to ovate-lanceolate,

sometimes slightly falcate, 10-20 cm long,

2-3-5 cm broad, smooth on upper surface,

with unevenly prominent veins on lower

surface, remotely 2-5-toothed on upper

margin towards base of leaflet, 1-3-toothed

on lower margin, with pungent apex. Cones

2-4 together on short stout tomentose

peduncles. Male cones subcylindric, 30-40

cm long, 13-15 cm diam., obtusely rounded

at base and apex; median scales 4-4-5 cm-

long, 2-5 cm broad with sharp lateral ridges;

bulla face projecting 1-1-5 cm, with deci-

duous reddish hairs, upper and lower facets

variably slightly angled, terminal facet 1 • 5-

1 -6 broad, 8-13 mm wide vertically. Female

cones oblong-oval in outline, 50-80 cm long,

20-30 cm diam.; median scales about 8 cm

long, 6-7 cm broad, 3-4 cm thick vertically;

bulla face projecting about 2 • 5 cm with fairly

acute lateral ridges extending into the in-

curved lateral lobes 1-2 cm long, slightly

verrucose to nearly smooth on exposed

portion, upper and lower facets rounded with

slight ridging, terminal facet thinly fawn-

tomentose, 1 • 5-2 cm broad, 1-1 • 5 cm wide

vertically. Seed red, angled by compression,

4-5-5 cm long, 2-5 cm diam., with fleshy

beak.

On mountains in the mist belt of the Soutpansberg

and Letaba districts, occurring in dense formation

at Modjadji’s Location. Map 4.

Transvaal. — Letaba: Modjadji Mt., Burtt Davy

H. 3086; Astley Maberly in PRE 15059; Verdoorn

& Dyer 2220 ; 2221; Compton 180891; Codd & De

Winter 3102; 3103; 3104; Rogers 22068. Soutpans-

berg: Codd 6918; Gerstner 5895; 6193; Verdoorn

& Dyer 2218; Key 1; 2; Compton 18061.

The Modjadji Cycad is the grandest species of

them all. The leaflets recurve from the place of

insertion on the rhachis and the teeth on the upper

margin of leaflets are often more conspicuous

towards the base than those on the lower margin.

Female cones weigh up to 75 lb. In FI. Cap. 5, 2

(Suppl.) 1933, Hutch. & Rattr. associated this

species with their broad concept of E. altensteinii.

21. Encephalartos paucidentatus Stapf &

Burtt Davy in Burtt Davy FI. Transv. 1 : 40,

99 fig. 4A (1926); Hutch. & Rattr. in F.C.

5, 2 (Suppl.) : 40 (1933); M. R. Henderson

in Journ. S. Afr. Bot. 11 ; 48 (1945); R. A.

Dyer in Bothalia 8 : 498 (1965). Type:

Transvaal; (exact locality and collector un-

certain) H. 5185 (PRE, holo.), see R. A. Dyer

in S. Afr. Journ. Sci. 61 : 50 (1965).

Plant unbranched or occasionally

branched from base, stems up to about 6 m

tall, 40-70 cm diam. with brown wool on

crown. Leaves 1-2 m long including petiole

20-40 cm long; rhachis usually fairly straight

or slightly recurved, sometimes slightly

twisted, pubescent at first, glabrescent with

age except woolly pulvinus; leaflets widely

spaced towards base, a few becoming smaller,

lobate and prickle-like, those towards apex

becoming narrower and closer together but

only sometimes overlapping; median leaflets

narrowly lanceolate, straight or somewhat

falcate, 15-25 cm long, 2-3-2 cm broad,

rarely more, with up to about 30 conspicuous

raised veins on lower surface, shortly pube-

scent, glabrescent on upper surface and more

slowly so on under surface, with or without

1- 3 prickles on one or both margins usually

towards base. Cones probably 2-3 together

on short stout peduncles, golden-yellow.

Male cones up to about 60 cm long, 15 cm

broad ; median scales narrowly oblong-

ovate to oblong-lanceolate, 4-5 cm long,

2- 2-5 cm broad, with sharp lateral ridges;

bulla projected into a decurved beak up to

about 1 ■ 5-2 cm long, upper and lower facets

humped towards base, terminal facet slightly

more than 1 cm broad and slightly narrower

vertically with irregularly toothed or crenate

margin. Female cones (complete cone not

seen) ; median scales 5-6 cm broad and about

3- 5 cm thick vertically with lateral ridges

extending into incurved lateral winged lobes

about 2 cm long; bulla face protruding

about 3 cm, with upper and lower facets

rounded and conspicuously rugose, apical

facet pubescent, concave, about 1 • 5-2 cm

broad and about the same width vertically,

with crenate margin. Seed red, up to about

4 cm long and 2 • 5 cm diam., with fleshy beak.

In association with low forest and mountain bush

in the eastern Transvaal and Swaziland, rare. Map 4 _

Transvaal. — Barberton: Van Elden in PRE

10085; Dyer 4800; Codd 9458; Thorncroft sub

Rogers 28426 (K); J. Thorncroft in PRE 29815.

Swaziland. — Miller 3597; Halker s.n.

The question of the locality from which the type

specimen was collected is discussed in the S. Afr.

Journ. Science, 61 : 50 (1965). The stated locality

north of the Soutpansberg is highly unlikely. Long

Zamiaceae

27

Fig. 7 — £. ferox, a, apical portion of leaf, reduced; b, apical portion of leaflet (taken trorn F.C. 5, 2 (Suppl)

' 1933).

28

Zamiaceae

slender tapering leaflets with conspicuous veins on

the lower surface are noteworthy characters of this

species.

22. Encephalartos ferox Bertol. f. in

Mem. Accad. Sci. Bologn. 3 : 264 (1851);

Prain in Kew Bull. 1916 : 180 (1916); F.T.A.

6, 2 : 352 (1917); Schuster in Pflanzenr. 4,

1 : 113 (1932); Lewis in F.Z. 1 : 81, t.l, fig. B

(1960); Paiva in Estud. Bot. 28 : 21 (1961);

R. A. Dyer in Bothalia 8 : 499 (1965). Type:

Mozambique, Fornasini.

E. kosiensis Hutch, in Kew Bull. 1932 : 512 (1932);

Hutch. & Rattr. in F.C. 5, 2 (Suppl.) : 34 (1933);

Compton in Hook. Icon, t.3220 (1933); Ogalvie in

Kew Bull. 1939 : 655, PI. 4 (1939); M. R. Henderson

in Journ. S. Afr. Bot. 11 : 27 (1954); Schelpe in

S. A. Journ. Sci. 47 : 16 (1950). Syntypes: Natal,

Zululand, Aitketi & Gale 63; Lugge NH 16507.

E. sp., Aitken & Gale in Bot. Survey S. Afr. Mem.

2 : 18 (1921).

Stems unbranched or occasionally

branched from base, majority less than 1 m tall

and only rarely are they 2 m or more tall,

about 30 cm diam. Leaves several to many in

a whorl, 1-2 m long including petiole 20-30

cm long ; rhachis nearly straight, erect spread-

ing, at first tomentose, glabrescent except for

pulvinus; leaflets mainly overlapping in the

upper half, more spaced in lower half, reduced

in size gradually towards base of rhachis into

a few to several prickles; median leaflets

broadest towards apex, more or less oblong-

elliptic, up to about 15 cm long and 3-5 cm

broad, rarely up to 5 cm broad, 2-4 small

teeth on upper and lower margin and termin-

ating in 3-5 broadly triangular pungent lobes

at the apex. Cones 1-3 together on stout

short peduncles, glabrous, shrimp-pink to red

(Brazil Red, R.C.S.). Male cones subcylindric,

narrowed to both ends, up to about 40 cm

long, 7-10 cm diam.; median scales 3-4 cm

long, 2-5-3 cm broad, 9-15 mm thick verti-

cally with sharp lateral ridges ; bulla face pro-

jecting into a decurved beak about 1 cm long,

upper and lower facets somewhat rugose,

terminal facet 6-10 mm broad, and about 6

mm wide vertically. Female cones ovoid to

ovate-oblong in outline, 25-50 cm long,

20-40 cm diam., median scales 5-6 • 7 cm long;

bulla 4-5-7 cm broad, 3 -5-5 -5 cm thick

vertically, with lateral ridges extending into

incurved lateral lobes almost to the main axis,

upper and lower facets finely wrinkled in basal

half, somewhat rugose on apical half, round-

ed, terminal facet slightly concave, 2-3 cm

broad and about 2 cm wide vertically. Seeds

scarlet, 4-5-5 cm long, 1 -5-2 cm diam., angled

by compression, with fleshy beak. Fig. 7.

Locally common in coastal bush and grassveld

from about 400 miles north of Lourenco Marques

to Sordwana Bay in Zululand; a solitary plant

reported about 25 miles inland from Sordwana Bay.

Map 4.

Natal. — Zululand, Aitken & Gale 63; Lugge

NH 16507; Prosser 1984.

The plant reported about 25 miles inland from

Sordwana Bay seems to be outside the normal range

of distribution and one can only speculate on how

it reached there.

23. Encephalartos villosus Lem., Illustr.

Hort. 14, Misc. 79 (1867); 15 : pl.557 (1868);

Dyer in Bot. Mag. t.6654 (1882); Marloth,

FI. S. Afr. 1 : 96, t.15 B, t.16 B (1913);

Schuster in Pflanzenr. 4, 1 : 1 18 (1932), in part,

excl. Zamia villosa Gaertn. ; Hutch. & Rattr.

in F.C. 5, 2 (Suppl.) : 31 (1933), excl. Z. villosa

Gaertn.; M. R. Henderson in Journ. S. Afr.

Bot. 11 : 17 (1945); Verdoorn in Flow. PI.

Afr. 26 : P1.1001, 1002 (1947); R. A. Dyer in

Bothalia 8 : 502 (1965). Type: Hort. Ver-

schaffelt, without locality, probably Natal, no

preserved specimen traced but tab. in Illustr.

Hort. 15, pi. 557 (1868) adequate; see R. A.

Dyer in Journ. S. Afr. Bot. 31 : 119 (1965).

E. mackenii sensu G. Henderson, 111. Diet. Gard.

1 : 508 (1885) not of Miq. Type: Natal, McKen,

Hort. ? E. striatus Stapf & Burtt Davy in Burtt

Davy, FI. Transv. 1 : 40, 98 (1926). Type: without

locality, Reid H.2987 (PRE 4, holo.).

Stem subterranean or sometimes with

the crown slightly exposed, usually unbranch-

ed, up to about 30 cm long, 20 cm diam. with

woolly interseriate bracts 5-7 cm long. Leaves

1-25-2-5 m long, suberect and gracefully

curved-spreading above, white-woolly on

rhachis and lower surface of leaflets when

young, glabrescent except for woolly pulvinus ;

leaflets glossy-green, spreading to recurving,

not overlapping except occasionally towards

apex of leaf, reduced to many prickles nearly

to base of rhachis; median leaflets linear to

linear-lanceolate, acuminate, 15-25 cm long

(rarely 30), 1-5-2 -5 cm broad, usually with

both upper and lower margins with 1-3 for-

ward-directed sharp prickles, rarely entire,

apex with single point or occasionally with 2

subequal lateral prickles in addition. Cones

1-several from a crown on stout peduncles

up to about 20 cm long and 2-5-5 cm diam..

Zamiaceae

29

c

Fig. 8.— E. villosus, a, base of leaf, reduced; b, tiplof leaf, reduced; c, median leaflet, reduced; d, tip of

leaflet (taken from F.C. 5, 2 (Suppl.) 1933).

30

Zamiaceae

greenish-yellow to orange-yellow. Male cones

up to about 65 cm tall, tapered to the apex

and 6-5-12 cm diam. above the base (varying

considerably and partly dependent on whether

one or more cones arise from one stem), thinly

pale brown tomentose when very young,

glabrescent, unpleasant smell when dehiscing;

scales set at upward angle from the axis;

median scales 3-5 cm long, 3-3-5 cm broad;

bulla laterally angled, with flattened face and

up to 1-5 cm thick vertically, upper facet

humped, terminal facet about 5 mm broad

and 5 mm wide vertically extending to the

lower margin of the face and there irregularly _

toothed or rough. Female cones oblong-ovoid

in outline, 30-40 cm long, 12-13 cm diam.;

median scales 4-4-5 cm long with quadrangu-

lar stipe about 2-6 cm long; bulla 4-5-5 cm

broad, 2-5 cm thick vertically with incurved

lateral lobes about 1-5 cm long and laterally

angled; bulla face flattened with the upper

and terminal facet almost in one plane; ter-

minal facet with the lower margin more or less

toothed and somewhat overlapping the scale

below. Seeds scarlet, 2 • 5-3 • 3 cm long, 1 • 8-2

cm diam., with fleshy apex. Fig. 8.

From the eastern Cape near East London to

Natal, south-eastern Transvaal and Swaziland, often

locally common and usually associated with bush or

low forest. Map 5.

Map 5. — Distribution map of E. villosus, E.

umbeluziensis, E. ngoyanus and E. cafTer.

Cape. — East London: Galpin 3340; 7102; 7767;

Dyer 4429. Kentani : Pegler 342; 756; 1124;

Flanagan 1374. Willowvale: Acocks 12287; Marais

485. Port St. Johns: Mogg 13052.

Natal. — Camperdown: Dver 4478; Edwards

2864. Zululand : BW4085; Wells 137.

Swaziland. — Stegi, Nicholson in PRE 27261;

Dyer 4794.

Transvaal. — Piet Retief: Du Plessis 114B;

Dyer & Verdoorn 5836.

Galpin’s reference on the label of one sheet of

his 3340 to a stem of 5 ft must have been the result

of confusion at some stage with E. altensteinii, with

which E. villosus sometimes grows in low forest near

East London. Specimens received at the Pretoria

National Botanic Garden from that area strongly

suggest that hybridization between these two species

takes place on occasions in the wild. Hybridization

between E. lebomboensis and E. villosus is indicated

in specimens seen inland from Pongola Poort in

the Piet Retief District. It seems that the farther

north one travels the more robust the form of this

species.

24. Encephalartos umbeluziensis R. A.

Dyer in Flow. PI. Afr. 28 : PI. 1100 (1951);

R. A. Dyer in Bothalia 8 : 505 (1965). Type:

Mozambique, Umbeluzi, Kev in PRE 28429

(PRE, holo.).

E. villosus sensu Lewis in F. Z. 1 : 83 (1960).

E. villosus var. umbeluziensis (R. A. Dyer) Lewis,

op. cit. 1 : 565 (1961); Paiva in Estud. Bot. 28 : 22

(1961). E. sp. aff. E. caffer Lehm. vel aff. E. villosus

Lem., M. R. Henderson in Journ. S. Afr. Bot. 11:17

(1945).

Stem subterranean or sometimes with

crown slightly exposed, usually unbranched

with thick tuberous root, up to about 30 cm

long and 20-25 cm diam., with densely woolly

interseriate bracts. Leaves 1-2 m long, in-

cluding petiole 10-30 cm long, suberect,

woolly at first except on upper surface of

leaflets, glabrescent; rhachis nearly straight,

sometimes spreading-recurved in shade, 7-12

mm thick towards base; leaflets light green

when young, soon becoming dark green,

spaced about 1 cm apart with the bases in-

serted on the rhachis more or less along a

straight line, twisted near the base like a half-

open Venetian-blind, reduced in size towards

base of rhachis but not reduced to a series of

prickles; median leaflets linear, tapering to

apex, 10-20 cm long, rarely upto 30 cm, 8-15

mm broad with apex pungent and usually

with 1-2 prickles on upper margin and 1-3 on

lower margin. Cones 1-3 together, peduncu-

late, olive-green turning to sulphur-yellow,

glabrous. Male cones subcylindric, tapering

slightly to apex, up to about 30 cm long, 6-8

cm diam. with peduncles 10-12 cm long, 2-5-

3 cm thick; median scales about 5 cm long,

2-2-4 cm broad, 1-1-2 cm thick vertically,

with acute lateral ridges; bulla face with the

upper facet humped or ridged, the lower

Zamiaceae

31

facet slightly rounded, terminal facet rhom-

boid, 8-12 mm broad and 6-9 mm wide ver-

tically. Female cones on peduncles about 15

cm long and 3 • 5 cm diam., up to 30 cm long,

12 cm diam.; median scales up to about 5 cm

long and broad and 3 cm thick vertically,

with acute lateral ridges; bulla face with

upper facet with 1-2 sagittal ridges, lower

facet slightly rounded and inconspicuous,

terminal facet rhomboid, 1-3-2 cm broad

and about the same vertically, somewhat

below centre of bulla face or occasionally

extending nearly to lower margin. Seed

scarlet, about 3-5 cm long, 2 cm thick, angled

by compression. Figs. 9a and 9b.

In dry forest without much undergrowth in

southern Mozambique and extending into Swaziland

up the Umbeluzi River Valley. Map 5.

Swaziland. — Compton 2957 1; 32066; 32067.

The basal leaflets are not reduced to a long series

of prickles as in E. villosus Lem. and the cones,

especially the male ones are considerably smaller

on the average. Young plants of E. umbeluziensis

and E. ngoyanus are not readily distinguished but so

far no overlap in distribution has been observed.

Mature plants of the former are considerably more

robust especially in the round petioles and larger

cones.

25. Encephalartos ngoyanus Verdoorn in

Flow. PI. Afr. 27: PI. 1053, 1054 (1949); R.

A. Dyer in Bothalia 8 : 508 ( 1 965). Syntypes :

Zululand, Verdoorn & Christian 716; 716b

(female and male respectively (PRE).

E. caffer sensu Hutch. & Rattr. in F.C. 5, 2

(Suppl.) : 30 (1933), partly, as to Rattray 1278.

E. sp. aff. E. caffer Lehrn., M. R. Henderson in

Journ. S. Afr. Bot. 11 : 16 (1945).

Stem subterranean or with crown ex-

posed, usually unbranched, up to about 30

cm long and 20 cm diam., with woolly bracts.

Leaves 3-7 in a whorl, 60-120 cm long in-

cluding slender petiole 10-30 cm long; rha-

chis erect or slightly recurving with silky and

woolly off-white tomentum, glabrescent ex-

cept for pulvinus; leaflets thinly pubescent

at first, glabrescent, reduced in size abruptly

near apex of leaf and gradually towards base

but rarely to a prickle; median leaflets linear-

lanceolate, pungent, 7-8 cm long, 9-11 mm

broad, occasionally in shade up to 12 cm

long and 1-5 cm broad, usually with 1-3

teeth on lower margin, rarely entire. Cones

solitary on short stout peduncle, pale olive-

green turning yellow. Male cones more or

less cylindric, 20-25 cm long, 4 • 5-6 cm diam. ;

Fig 9a. — E. umbeluziensis, a, basal leaflets, not reduced to spines; c, tip of leaf, both X £ ( Dyer 4798).

Zamiaceae

32

Fig. 9b. — E. umbeluziensis, b, median leaflets; d, single median leaflet, both X \ ( Dyer 4798).

median scales 2 • 5-2 • 8 cm long, 2 • 2-2 • 5 cm

broad, 1-4-1 -5 cm thick vertically, laterally

angled ; bulla face projecting in a subcylindric

beak 7-8 mm long with the terminal facet

only faintly indicated except for lower margin

which is acute, decurved andsometimes slight-

ly overlapping scale below. Female cones

ovate-oblong in outline about 23 cm long,

10 cm diam. ; median scales about 5 cm long,

4-5 cm broad and 3 cm thick vertically,

laterally ridged and extended into incurved

lobes about 5 mm long; bulla face only

slightly projecting with upper facet humped,

terminal facet faintly defined on upper mar-

gin, lower margin acute decurved and some-

times slightly overlapping scale below. Seed

scarlet, 2-7-3 cm long, 2 cm diam.

In grassveld and forest margins, often near

boulders, in northern Zululand and over the Trans-

vaal and Swaziland border. Map 5.

Natal. — Mtunzini: Ngoye, Verdoorn & Christian

715; 715a; 716; 716a, b, c; Rattray 1278; Ward

3469. Lower Umfolozi: Mkuzi, Ga/pin 13310.

Ubombo: Ward 4077. Ingwavuma: Verdoorn &

Christian 720; 720a, b, c, d; Conyngham la, b, c.

Transvaal.— Piet Retief: Pongola Poort, Wil-

liams in PRE 29821.

Swaziland. — Ubombo Mt., Ga/pin 11818.

Zamiaceae

33

This species seems to be distinguishable from E.

caffer in a number of inconspicuous ways. Firstly

the root-system is less tuberous, the stem less con-

sistently subterranean, the leaflets less crowded, not

twisted and the lower margin often toothed, and the

female cone scales have a less pronounced terminal

facet

The break in distribution between the two species

is noteworthy but not necessarily taxonomically

significant.

26. Encephalartos caffer ( Thunb .) Lehm.,

Pugill. 6 : 14 (1834); DC., Prod. 16, 2 : 532

(1868); Schuster in Pflanzenr. 4, 1 : 109

(1932); Hutch. & Rattr. F.C. 5, 2 (Suppl.):

29 (1933), excluding specimens cited from

Natal; M. R. Henderson, in Journ. S. Afr.

Bot. 11:13 (1945); R. A. Dyer in Bothalia

8:509(1965). Syntypes: Cape, Thunb. sheets

1 & 2 of leaves and part of male and female

cone material (UPS).

Cycas caffra Thunb., in Nov. Act. Soc. Sc. Uppsal.

2 : 285, (1775) partly, as to t.5 and part of Thunberg’s

material at Uppsala, excluding part of description.

Zamia cycadis L. f., Suppl. 443 (1781). Type:

Cape, Thunberg. Z. villosa Gaertn. Fruct. 1, t.3

(1788). Z. pimgens Ait. Hort. Kew ed. 1, 3 : 478

(1789) partly, as to Masson’s specimen. Z. caffra

Thunb.. Prodr. 2 : 92 (1800); FI. Cap. ed. Schult.

429 (1823) partly, as to Thunberg’s material and

Fig. of Cycas caffra.

Encephalartos brachyphyllus Lehm., Cat. Hort.

Hamb. 1836, ex Lehm. & De Vriese, Tydsch. Nat.

Gesch. 4 : 414, t.6 & 7 (1838) and Nov. Sp. Cvcad.

6, t.l & 2 (Scrip. 1837). Type: Hort. E. cycadis (L.f.)

Sweet. Hort. Brit. ed. 3 : 626 (1839). E. verschaffeltii

Regel in Act. Hort. Petrop. 3 : 3 (1875). Type: Hort.

E. caffrorum Hort. Sion ex Yates in Proc. Linn. Soc.

11:19 (1849); Schuster l.c. Type: Hort.

Plant with subterranean stem or with

crown slightly exposed; roots tuberous;

stem unbranched or occasionally branched,

up to about 30 cm long, 15-25 cm diam.,

woolly at crown. Leaves 40-90 cm long,

including the petiole of a few-several cm

long; rhachis erect at first or slightly curved,

densely brown woolly, glabrescent except

pulvinus; leaflets numerous and crowded,

sometimes twisted at various angles to the

axis, glabrous, reduced at base to 1-2

prickles; median leaflets linear-lanceolate,

up to about 10 cm long, 8-10 mm broad,

entire, although toothed on young plants,

somewhat decurrent at base. Cones solitary

on stout peduncles, glabrous, usually shining,

greenish-yellow. Male cones with peduncle

up to about 15 cm long, cylindric 20-30 cm

long, 6-11 cm diam., median scale 2 -5-3 -5

cm long, 2-5 cm broad with lateral ridges;

bulla face projecting 5-6 mm beyond spor-

angia, upper facet smooth rounded, terminal

facet 1-1-5 cm broad and wide vertically,

slightly below central, concave and with the

lower margin sometimes slightly toothed.

Female cones on peduncle 6-7 cm long and

3-5-4 cm diam., subcylindric, slightly nar-

rowed to rounded apex, about 30 cm long,

15 cm broad; median scales 4-5-6 cm long,

5 • 5-6 cm broad, 3 • 5-3 • 7 cm thick vertically ;

bulla with lateral ridges extending into in-

curved lateral lobes about 1 cm long, basal

portion pale orange-yellow or orange, closely

papillate, upper facet rounded, with or

without a sagittal ridge and fairly smooth

surface, lower facet rounded, terminal facet

2 • 25-2 • 5 cm broad, 1 • 5-2 cm wide vertically,

with slightly raised and sometimes toothed

margin. Seeds scarlet, 3 -7-3 -9 cm long,

2 -3-2 -5 cm diam.

In sour grassveld of coastal belt from southern

Steytlerville and Uitenhage Districts to the neigh-

bourhood of Willowvale in the Transkei ; infrequent.

Map 5.

Cape. — Steytlerville: cult. NGB 2601/16; Hender-

son 1505. Uitenhage: Van Stadens, Rattray 1098.

Alexandria: Archibald 4565. Bathurst: Verdoorn &

Christian 701; Dyer 4839; Smith 6558; Dyer &

Wells 5795. East London: Mt. Coke, Galpin 7839;

Smith 6901; Dyer 4845. Kentani: Pegler 1124;

1127; 2156; Verdoorn & Christian 710.

The species has not been rediscovered in the

Steytlerville or Uitenhage districts in recent years.

For many years there was uncertainty about the

correct application of the epithet caffer because

Thunberg confused two species under the one name.

The other species is now named E. longifolius Lehm.

G. G. Smith notes that specimens from the neigh-

bourhood of East London have less rigid leaves than

those from Bathurst and the leaflets are rolled

inwards in the young stage. M. Wells noted inrolled

young leaflets on a plant in Bathurst. On the other

hand plants from Zululand, formerly included in

E. caffer, are recognised as specifically distinct and

are to be found under the name E. ngoyanus.

While these notes were in press, a record of

E. caffer in the Humansdorp District has been

confirmed. It is likely that Thunberg collected his

material of both E. caffer and E. longifolius (which

he confused) in this district.

Doubtful or Excluded Species

The following names have been published but the

information is too incomplete for specific identifica-

tion. :

E. mauritianus Miq., Monogr. 48 (1842). This is

cited under E. lehmannii in FI. Cap. 5, 2

(Suppl.) 1933.

34

Zamiaceae

E. nanus Lehm. in Tydschr. 4 : 421 (1837). Related

to E. horridus Lehm. but not possible to

identify with certainty.

E. plumosus G. Henderson, 111. Diet. Gard. 508

(1885). This could refer to a form of E. villosus.

E. pungens (Ait.) Lehm. Pugill. 6:13 (1834 ) = Zamia

pungens Ait. Name based on a mixture of con-

cepts.

E. regalis G. Henderson, 111. Diet. Gard. Suppl. 341

(1900). This is thought to refer to a form of

E. altensteinii.

E. tridentatus (Willd.) Lehm. Pugill. 6.13 (1834).

= Zamia tridentata Willd. Sp. PI. 4 : 846 (1806).

Apparently based on a juvenile specimen of

uncertain origin.

E. van-hallii de Vriese in Hoev. & de Vriese, Tydschr.

4 : 422 (1837-38). This is cited under E.

latifrons in FI. Cap. 5, 2 (Suppl.) 1933 and may-

well have been correctly diagnosed.

E. vromii Matte., Rech. App. Lib. Lign. Cycad. 70,

(nomen) = £. altensteinii var. vromii G. Hender-

son, 111. Diet. Gard. 508 (1885).

The following epithets and others cited by Schuster

in Pflanzenr. Cycad. (1932) which we have been

unable to verify do not appear to have nomencla-

tural significance:

E. almasianius Hort. under E. longifolius

E. aquifolius Lodd. under E. horridus var. latifrons

E. brownei Hort. under E. horridus var. latifrons

E. crassifolius Hort. under E. cajfer

E. ellipticus Lodd. ex Miq. under E. caffer

E. glaber Hort. under E. altensteinii

E. grandis Hort. under E. altensteinii var. eriocephalus

E. katzeri Regel ex Schuster under E. altensteinii

E. katzeri Hort. under E. horridus var. latifrons

E. lepeschkinei hort. under E. horridus

E. macrophyllus Hort. under E. horridus

E. niveo-lanuginosus Wendl. in Hort. under E.

villosus

E. procer Miq. under E. elongatus

E. pumilus Lehm. MS. under E. horridus var. nanus

E. revolutus Hort. under E. longifolius

E. royeni Miq. in Herb, under E. caffer

E. van-den-heckei Hort. under E. altensteinii var.

eriocephalus

E. van-geertii Miq. under E. cycadifolius var.

friderici-guilielmi

PODOCARPACEAE

by O. A. Leistner

Trees or shrubs, usually dioecious. Leaves linear to ovate or scale-like. Male cones

axillary or terminal, usually consisting of numerous fertile scales each bearing 2 pollen sacs;

pollen grains with air-bladders or wings. Female cones reduced, with 1 to several fertile scales;

ovule solitary, erect or inverted, usually enclosed in a false aril (epimatium). Cotyledons 2.

Genera 7, found chiefly in the tropics and the southern temperate zone, in Africa, Asia, Australasia and

South America. Only one genus occurs in Southern Africa.

In Phillips’s Genera the genus is placed under the Taxaceae, but in agreement with Pilger (Engl. & Prantl,

Pflanzenf. ed. 2, 13 : 211, 1926) and subsequent authors such as Stapf (F. C. 5, 2 Suppl.: 3, 1933) and

Florin (Acta Hort. Berg. 15: 285-388, 1951) it is here put into a separate family.

In accordance with a recommendation of the International Code of Botanical Nomenclature the name

Podocarpus is here treated as masculine. Previously it has, however, often been regarded as feminine and

where such names are quoted under synonymy in the present work they are given in the original form.

13 PODOCARPUS

Podocarpus UHerit. ex Pers., Syn. 2 : 580 (1807), nom. cons.; Endl., Gen. PI. 1 : 262 (1837);

Pilg. in Pflanzenr. 4, 5 : 54 (1903); Stapf in F.C. 5, 2 (Suppl.) : 3 (1933); Buchholz & Gray

in Journ. Arn. Arb. 29 : 54 (1948); Phill., Gen. ed. 2 : 50 (1951).

Nageia Gaertn., Fruct. 1 : 191 (1788), partly, nom. rej.

Trees or shrubs, dioecious, evergreen. Leaves spirally arranged, subopposite or whorled,

rarely opposite, linear to narrowly ovate or scale-like. Male cones axillary, rarely terminal,

sessile to shortly stalked, catkin-like; pollen grains usually with 2 air-bladders or wings.

PODOCARPACEAE

35

Female cones axillary, on thin naked stalks or on short leafy or scaly branches, usually reduced

to 1 to 2 fertile terminal scales and a few sterile lower scales which are fused with each other

and with the cone axis; the fused cone axis and scales beneath the seeds either remain woody

or become swollen and often fleshy at maturity (receptacle). Ovule solitary, adnate to the face

of the fertile scale, inverted and enclosed in a false aril (epimatium) arising from the face of

the scale and adnate to the single integument. Seeds ellipsoid to globose; testa and false aril

form a coriaceous or externally fleshy and internally more or less woody shell; embryo axile.

Found mainly in montane and coastal forests in the tropics and the southern subtropics. There are about

80 species in the genus, of which four are recognised in Southern Africa.

The shape of leaves of all species varies greatly depending on the age of the specimen and the position of

the leaf on the shoot. In general, leaves of juvenile specimens are longer, more slender, more pointed and

often more falcate than leaves of adult trees. Juvenile features are usually lost when the tree is about 5 to 10

years old but they are sometimes retained in specimens of appreciable age. Leaves of senile plants tend to

differ as follows from those of normal adult specimens: they are shorter, and often either narrower (e.g.

P. falcatus) or wider (e.g. P. latifolius).

In both juvenile and adult specimens of all species dealt with, the 2 or 3 lowermost, and thus oldest,

leaves on each shoot differ as follows from average leaves: they are shorter, broader, blunter, tend to be

widest in the upper half and often have a brown tip. The uppermost 2 or 3 leaves, on the other hand, are

generally longer, narrower and more acute than average leaves and they tend to be widest in the lower half.

This variability must be considered when consulting the key and the descriptions, which are based largely

on leaves from the middle of shoots of average adult specimens.

Leaves of all species in our area are frequently infested by a fungus, Corynelia uberata Fr. ex Ach.

Some of the specimens cited in the following references were not seen: Pilg. in Ptlanzenr. 4, 5 (1903),

Burtt Davy, FI. Transv. (1926), Stapf in F.C. 5,2 (Suppl.) (1933), Gray in Journ. Arn. Arb. 34 (1953).

Branchlets usually less than 1 -5 mm in diameter near apex, often square and distinctly ridged by leaf

bases; leaves twisted at base and blade thus orientated in vertical plane, stomata present on both

surfaces; seeds borne on scaly or leafy branches, which do not become swollen at maturity;

Swellendam eastward into Transvaal 1 . P. falcatus

Branchlets usually more than 1-5 mm in diameter near apex, terete, with shallow grooves or slight

ridges; leaves not twisted at base and blade thus orientated in horizontal plane, stomata absent or

sparse on upper surface; seeds borne on receptacles which become swollen and often fleshy at

maturity, receptacles on naked stalks:

Leaves usually shorter than 8 cm, roughly parallel-sided in middle portion, spreading to erect; seeds

borne on succulent, often brightly coloured receptacles:

Leaves generally wider than 5 mm, stomata absent on upper surface (very rarely a few present);

Cape Peninsula and eastward into Transvaal 2. P. latifolius

Leaves generally 4-5 mm wide, many with 1 to several short rows of stomata on upper surface;

western Cape only 3. P. elongatus

Leaves usually longer than 8 cm, tapering gradually from about the middle to an acuminate tip, more

or less pendulous; seeds borne on receptacles which are slightly swollen but neither succulent nor

brightly coloured; Natal, eastern Cape 4. P. Iienkelii

1. Podocarpus falcatus ( Thunb .) R.Br. ex

Mirb. in Mem. Mus. Hist. Nat. Paris 13 : 75

(1825); Endl., Syn. Conif. 219 (1847);

Carriere, Traite Conif. ed. 1 : 472 (1855);

Henkel & Hochstetter, Syn. Nadelholz. 400

(1865); Pari, in DC., Prodr. 16,2:511 (1868);

Pilg. in Pflanzenr. 4, 5 : 72 (1903), not seen:

Ecklon & Zeyher s.n., Bachmann 74, Drege

6182, Bachmann 69; Burtt Davy, FI. Transv.

1 : 101 (1926), partly, excl. Brit. Bech., nr.

Genesa ; Chalk et al.. Forest Trees & Timbers

Brit. Emp. 1 : 23 (1932); l.c. 3 : 86 (1935);

Marloth, FI. S. Afr. 3, 1 : 66, t.19 (1932);

Stapf in F.C. 5, 2 (Suppl.): 10 (1933), partly,

excl. Wood 3005 and Brit. Bech.: Stellaland,

not seen: F.D. Herb. 1246, 1244, Bachmann

74, Sanderson 3015, Rehmann 6482; Robyns

in Bull. Inst. Roy. Col. Belg. 6 : 237 (1935);

Dallimore & Jackson, Handb. Conif. 67

(1948); Gray in Journ. Arn. Arb. 34 : 69

(1953), probably excl. Burtt Davy 20248, not

seen: Anon s.n. ex Scheidweiler Herb., Basil

s.n.. Anon s.n. ex Herb. A.N.S., Anon s.n.,

Molliana s.n., H.S. s.n., “Hort. Daudin”,

“Hort. Monac.”, Engelmann s.n., Whitford

27; Melville in Kew Bull. 1954: 568 (1955);

Lewis in F.Z. 1 : 85 (1960). Type: Ribeck-

castel, Vleermuysdrift, Thunberg 23779 (UPS,

holo. !).

36

PODOCARPACEAE

Taxus falcata Thunb., Prodr. 117 (1800); FI.

Cap. ed. Schultes 547 (1823).

Podocarpus meyeriana Endl., Syn. Conif. 218

(1847); Pari, in DC., Prodr. 16, 2 : 512 (1868).

Type: Drege s.n. (?) P. elongata sensu Pappe,

Silva Capensis 32 (1854); Carriere, Traite Conif.

ed. 2 : 670 (1867), partly; Sim, For. FI. Cape Col.

335 (1907); Marloth, FI. S. Afr. 1 : 101 (1913),

non (Ait.) L’Herit. ex Pers. P. falcatus var. lati-

folius Pilg. in Pflanzenr. 4, 5 : 72 (1903). Type:

Kaymansgat, Drege 6182 (B, holo.). — var. pon-

doensis Pilg. in Pflanzenr. 4, 5 : 73 (1903). Type:

Pondoland, Bachmann 69 (B, holo.). P. thunbergii

var. falcata (Thunb.) Sim, For. FI. Cape Col. 332

(1907), partly, as to name and synonymy but excl.

description. P. gracilior sensu Burtt Davy, FI.

Transv. 1 : 101 (1926); Stapf in F.C. 5, 2 (Suppl.):

13 (1933). P. gracillimus Stapf in F.T.A. 6, 2 : 343

(1917); Burtt Davy, FI. Transv. 1 : 101 (1926);

Chalk et al.. Forest Trees & Timbers Brit. Emp.

1 : 23 (1932); Stapf in F.C. 5, 2 (Suppl.): 14 (1933);

Gray in Journ. Arn. Arb. 34 : 73 (1953). Type:

Houtboschberg, Nelson 423 (K, holo.!).

Tall tree generally 10-25 m high but

attaining a height of 60 m with a clean bole of

more than 20 m and a girth of about 7 m.

Bark greyish to purplish, more or less smooth

and persistent in young trees, flaking in

rectangular to roundish pieces in older speci-

mens. Branchlets terete or square (on juvenile

specimens generally square), distinctly ridged

by decurrent leaf bases. Terminal buds about

1 mm in diameter; outer bud scales very

narrowly triangular, 2-2-5 mm long and

about 1 mm wide. Leaves spirally arranged,

on branchlets of juvenile specimens often

subopposite, spreading to suberect, glaucous

to yellowish-green, twisted at the base and

lamina thus orientated in a more or less verti-

cal plane, narrowly linear-lanceolate to linear-

elliptic, falcate to straight, acute to obtuse;

adult leaves (1-) 2-4 (-4-5) cm long and

(1-2-) 2-4 (-6) mm wide; juvenile leaves up

to 12 cm long and 0-6 cm wide; midrib

slightly raised on lower surface, very slightly

raised on upper surface; stomata present on

both surfaces, arranged in 14-20 ± distinct

longitudinal lines on either side of midrib.

Male cones solitary or in groups of 2-4,

subsessile to very shortly stalked, 5-13 mm

long, elongating up to 15 mm after shedding

pollen, (2-) 3 (-3 • 5) mm in diameter, brown-

ish ; outer sterile scales at base very broadly

triangular-trullate to very broadly obovate,

crenulate to denticulate, 0-5-1 mm long and

1-1-5 mm wide; terminal lobe of fertile

scales very broadly triangular-trullate,

0-6-0 -8 mm long and 0-8-1 -4 mm wide,

crenulate to lacerate; pollen sacs 0-6-0 -7

mm long and about 0-3-0 -4 mm in diameter.

Female cones solitary on scaly or leafy

branches 7-27 mm long and 1 • 5-2 • 5 mm in

diameter, widest at the top just below seed;

only the terminal scale fertile. Seed sub-

spherical to obovoid, ( 1 - 2—) 1-3-1 -7 (-1-8)

cm long, glaucous to greyish-green, ripening

to a yellowish or light reddish-brown colour;

testa consisting of outer somewhat fleshy

covering up to 3 mm thick which becomes

very resinous inwards, and inside this a sub-

spherical, somewhat laterally compressed

tubercled kernel 1-1-2 (-1-4) cm long with

hard woody walls (0 • 8-) 1-1 • 7 (-2) mm thick.

Fig. 10 : 1.

This, the tallest member of the genus in Southern

Africa, occurs in coastal and montane forests from

the Swellendam district in the Cape to the northern

Transvaal and southern Mozambique. It is much

less common than P. latifolius and apparently is

only rarely dominant.

Cape. — Albany: near Grahamstown, Jacot

Guillarmod 4600 (RUH). Alexandria: Langebosch,

Archibald 4499. Bathurst: Kariega River, Britten

2440. East London: Marloth 2587. Engcobo:

Buswayo Forest, Van der Merwe in F.D. Herb.

2268. George: Wilderness, Marloth 12703. Humans-

dorp: Scott’s Cave, Wells 2851. Kentani: near

Kentani, Marais 739. King William’s Town: near

King William’s Town, Comins 1027 (GRA). Knysna:

Plettenberg Bay, Rogers 26861. Komga: near

Komga, Flanagan 1179. Lusikisiki: Ntsubane

Forest, Fraser in F.D. Herb. 2226. Mossel Bay:

above Langfontein, Muir 2380 (J). Mount Ayliff:

Fort Donald Forest Station, Cochrane s.n. Port

Elizabeth: Van Staden’s River, Theron 1658.

Queenstown: Bongolo Nek, Galpin 7973. Somerset

East: Boschberg, Burchell 3174 (K). Stocken-

stroom: Katberg, Moss 15339 (J). Stutterheim:

Kabaku Hills, Acocks 8998. Swellendam: Groot-

vadersbosch, Marloth 3496. Uitenhage: Kamachs,

Long 1280.

Natal. — Bergville: Indumeni Forest, Killick &

Marais 2142. Estcourt: Balgowan, Hilliard 179

(NU). Helpmekaar: near Pomeroy, Hilliard 1569

(NU). Ixopo: Ingwangwane, Houshold in F.D.

Herb. 1956. Ingwavuma: Kosi Bay, Edwards 2551.

Ladysmith: Cundycleugh, Hilliard 1031 A (NU).

Lions River: Karkloof, Hilliard 2026 (NU). Polela:

near Donnybrook, Hilliard 2464 (NU). Umfolozi:

near Kwambonambi Halt, Ballenden in F.D. Herb.

2925. Utrecht : Donkerhoek, Devenish 665. Weenen:

Umhlumba Mountains, West & Acocks 2745 (NH).

Swaziland. — Stegi: Ubombo Mountains, Miller

S/19.

Transvaal.- — Barberton: Ameide Plantation,

Scheepers 1242; 1243. Letaba: near Ofcolaco,

Scheepers 1241. Pietersburg: near Haenertsburg,

Codd 9437. Pilgrims Rest: Mariepskop, Van der

Schijff 5142. Soutpansberg: Houshold in Col. Herb.

5248.

PODOCARPACEAE

37

Fig. 10. — 1, Podocarpus falcatus, portion of branch with seed ( Watsham s.n.); la, portion of branch with

male cones (Jacot Guillarmod s.n.). 2, P. latifolius, portion of branch with mature receptacle and seed

( Von Breitenbach s.n. in George For. Herb. 1004); 2a, portion of branch with male cones (Von Breiten-

bach s.n. in George For. Herb. 1002). 3, P. elongatus, portion of branch with mature receptacles and

seeds (Kerfoot s.n.); 3a, male cone (Garside 1044). 4, P. henkelii, portion of branch with seed ( Bird s.n.

in George For. Herb. 1001); 4a, male cone (Von Breitenbach s.n. in George For. Herb. 1005).

38

PODOCARPACEAE

In some specimens from the Transvaal the leaf

shape approaches that of the tropical African species

P. gracilior Pilg., i.e. the leaves are more slender

than is usual in those of P. falcatus and they taper

gradually towards the tip from a point about the

middle of the leaf, while those of P. falcatus normally

tend to be more parallel-sided. Stapf regarded

these slender-leaved specimens as true P. gracilior.

A few Transvaal specimens, in which the leaves are

markedly reduced in size, Stapf placed into a

separate species, P. gracillimus. As regards their

reproductive structures none of the Transvaal

specimens were found to differ significantly from

the typical P. falcatus and they are thus regarded

as belonging to the same species. Specimens from

the Ingwavuma district in northern Natal differ

from the typical form in having leaves which are

wider and more nearly parallel-sided, and seeds-

which are often larger and have thicker woody walls.

The record from British Bechuanaland in F.C.

5,2 (Suppl.): 12 (1933) refers to Terminalia sericea

Burch, ex DC. which is commonly known as

“Geelhout” ( = Yellowwood).

Probably the tallest tree in South African forests;

it yields valuable timber provided the wood is well

seasoned. Monkeys and birds are reported to eat

the fruit. Some vernacular names are: Outeniqua

Yellowwood, Bastard Yellowwood, Outeniekwa-

geelhout and umSonti.

2. Podocarpus latifolius ( Thunb .) R.Br. ex

Mirb. in Mem. Mus. Hist. Nat. Paris 13 : 75

(1825), non Wall.', Pilger in Pflanzenr. 4, 5:

90. fig. 17 B (1903), partly, excl. MacOwan

1958 (1598 sphalrn.) in part, Gotze 986, not

seen: Ecklon & Zeyher 259, Beyrich 92, 163,

Bachmann 70, 71, 72, Gotze 986, Bergius s.n.,

Schlechter 3947; Marloth, FI. S. Afr. 1 : 101,

102, 1. 1 3, 1. 1 7 A (1913); Burtt Davy, FI.

Transv. 1 : 100, fig. 5 : A-D (1926), not seen:

Davy 2467, 2748. 5114, 5146, Legal 2396 b, c,

d, 2397, H 2453, Rogers 2455, 21607; Chalk

et al., Forest Trees & Timbers Brit. Emp.

1 : 23 (1932); 3:81 (1935), partly, excluding

Gillett s.n. in FHO 30023; Stapf in F.C. 5, 2

(Suppl.): 5 (1933), partly, excl. MacOwan

1958 in part, Zeyher 3884, not seen : Sim s.n.,

Beyrich s.n., Fourcade s.n., Burtt Davy 2313,

Botha s.n.; Robyns in Bull. Inst. Roy. Col.

Belg. 6 : 240 (1935); Dallimore & Jackson,

Handb. Conif. 72 (1948); Adamson in FI.

Cape Penins. 31 (1950); Gray in Journ. Arn.

Arb. 34 : 167 (1953), partly, excl. F.D. Herb.

2170, Gilbert 505, Zeyher 3880, 3885, not seen:

Bowie s.n., Ecklon & Zeyher s.n., H.R.P. s.n.,

Eames s.n., Compton s.n., Cook s.n., Buchholz

s.n.; Lewis in F.Z. 1 : 86 (I960). Type:

Houtniquas, Grootvadersbosch, aliis, Thun-

berg 23780 (UPS, holo.!).

Taxus latifolia Thunb., Prodr. 117 (1800); FI.

Cap. ed. Schultes 547 (1823).

Nageia latifolia Kuntze, Rev. Gen. 2 : 800 (1891),

non Gord.

Podocarpus thunbergii Hook, in Lond. Journ.

Bot. 1 : 657, t. 22 (1842); Endl., Syn. Conif. 217

(1847); Pappe, Silva Capensis 32 (1854); Carriere,

Traite Conif. ed. 2 : 670 (1867); Pari, in DC.,

Prodr. 16, 2 : 511 (1868); Marloth, Kapland 190,

191 (1908). Type: Houtniquas, Grootvadersbosch,

aliis, Thunberg 23780 (UPS, holo.!). — var . falcata

sensu Sim, For. FI. Cape Col. (1907), partly, as to

t.149 fig. 2 only. — var. latifolia (Thunb.) Sim,

For. FI. Cape Col. 332, t.148 (1907). P. latifolius

var. confertus Pilg. in Pflanzenr. 4, 5 : 90 (1903).

Syntypes: Table Mountain, Cape Town, Bergius

s.n. and Schlechter 3947. — var. latior Pilg. in

Pflanzenr. 4, 5 : 90 (1903); Chalk et ah. Forest

Trees & Timbers Brit. Emp. 3 : 82 (1935); Gray in

Journ. Arn. Arb. 34 : 169 (1953). Type: Vogelgat

Mountains, Schlechter 9542 (K, iso.!). P. henkelii

sensu Burtt Davy, FI. Transv. 1 : 101 (1926), excl.

fig. 5G; Stapf in F.C. 5, 2 (Suppl.): 9 (1933),

partly as to Cooper 1298, Sim 19019, Burtt Davy

2748a (2738a sphalrn.), Legat 2467 (3467 sphalrn.).

Nelson 420; Gray in Journ. Arn. Arb. 34 : 169

(1953), partly as to Legat 2467 ( Barbarton 2467

sphalrn.) and Burtt Davy 2748a.

Tree up to 33 m high with straight clean

bole up to 3 m in diameter or stunted tree or

shrub no more than 2 m high. Bark dark grey

to khaki-coloured, smooth in young trees,

later longitudinally fissured and exfoliating in

long strips. Branchlets slightly angular,

marked with grooves from decurrent leaf

bases. Terminal buds 2-3 -5 mm in diameter;

outer bud scales narrowly triangular, acumi-

nate, upper half sometimes recurved. 4-6 mm

long and about 1 • 5 mm wide. Leaves spirally

arranged to subopposite, often crowded in

upper parts of shoots, spreading, dark green

to glaucous and shiny above; adult leaves

straight to very slightly falcate, linear-elliptic,

more or less parallel-sided in middle portion,

rather abruptly contracted in upper 1/4 or 1/5,

apiculate to subobtuse, (2-) 3-5-6 (-10) cm

long and (0-5-) 0-6-0 -8 (-1-3) cm wide;

juvenile leaves up to 17 cm long and up to 17

times longer than broad; midrib raised on

lower surface, on upper surface slightly raised

in lower 2/3; margins slightly reflexed;

stomata confined to lower surface (very rarely

a few present on upper surface), arranged in

20-35 ± distinct longitudinal rows on either

side of midrib. Male cones solitary, rarely in

groups of 2, sessile or subsessile, (0-8-) 1--2

(-3) cm long, elongating to 3 cm or more after

shedding pollen, (2-5-) 3-4-5 (-6) mm in

diameter, pinkish ; outer sterile scales at base

narrowly to very narrowly triangular, often

PODOCARPACEAE

39

distinctly keeled, subentire to denticulate,

3-4 mm long and about 1-5-2 mm wide;

terminal lobe of fertile scale broadly triangu-

lar to widely ovate, about 0-6 mm long and

0 • 6-0 -7 mm wide, lacerate ; pollen sacs about

1-3 mm long and 0-6-0 -7 mm in diameter.

Female cones solitary, on naked stalks (1 -5-)

5-10 (-19) mm long and about 1 mm in

diameter; receptacle fleshy, glaucous green,

turning pink to reddish-purple when ripe,

sweet-tasting and semi-transparent, 8-14 mm

long and 8-14 mm wide, with 1, less com-

monly 2 fertile scales, 1 or sometimes 2 seeds

maturing on each receptacle. Seed obovoid to

subglobose, often slightly apiculate, glaucous

to slate-coloured, sometimes turning dark

violet, 7-1 1 mm long ; shell 0 • 5-1 ■ 2 mm thick,

consisting of 3 layers : the outermost thin and

leathery, the middle one varies in thickness,

is slightly woody, often contains cavities filled

with resin, the inner thin and pergament-

aceous. Fig. 10 : 2.

Found in the Cape Peninsula, the high forests in

the southern districts of the Cape Province and

in coastal and midland climax forests from the

Eastern Cape to the northern Transvaal. Occurs as

a tall straight tree in high temperate forests, and as

a low spreading tree or shrub on exposed rocky

slopes and in open coastal bush.

Cape. — Albany: Paradise Kloof, Jacot Guillarmod

s.n. (RUH). Bredasdorp: Potberg, Pi/Ians 9454

(BOL). Cape: Houtbay, Marloth 1795. Caledon:

Vogelgat Mountains, Schlechter 9542 (K). Engcobo :

Mkonto Forest, Miller 3082. George: Mund s.n.

Flumansdorp: Assegaaibos, Thode A2574. Keis-

kammahoek: Ghulukop, Wells 3091 A.KingWilliam’s

Town: near King William’s Town, Comins 1026.

Knysna: Harkerville Forest, Keet 524. Komga:

near Komga, Flanagan 1728. Lusikisiki: Ntsubane

Forest, Fraser in F.D. Elerb. 2227. Maclear: Pot

River Berg, Galpin 6831. Mqanduli: Koffiebaai,

Van der Schijff 5446. Mount Ayliff: Gxwaleni

Forest, Cochrane in F.D. Herb. 2173. Port Eliza-

beth: Loerie, Dix 191 (GRA). Riversdale: above

Corente River, Muir 394. Stockenstroom: Katberg,

Gilliland in J 22532. Stutterheim: Kabaku Hills,

Acocks 8996. Swellendam: Duiwelsbos, Zeyher

3882. Tabankulu: Tabankulu Forest, FHUiard 2500

(NU). Umzimkulu: Mhlonga Forest, Kaufmann

in F.D. Herb. 2168, 2169.

O.F.S. — Harrismith: Retief Klip, Steytler in

NBG 56065; Swinburne, Boschkloof, Jacohsz 152.

Natal. — Alfred: Ngeli Slopes, Hilliard 1789

(NU). Bergville: Cathedral Peak, Killick 1645.

Durban: Berea Hill, Thode in STE 8044. Eshowe:

Gerstner 3107. Estcourt: Tabamhlope, 1 Vest 87.

Ixopo: Ingwangwane, Houshold in F.D. Herb. 1957

Kranskop: Ntunjambili, Hilliard 1433 (NU).

Ladysmith: Van Reenen, Hilliard 1034A (NU).

Lions River: Karkloof, Hilliard 2024 (NU).

Mtunzini: Ngoya Forest, Huntley 270. Newcastle:

Amajuba Hill, Phillips 76. Ngotshe: Tendeka,

Gerstner 5167. Nkandla: Qudeni, De Winter 8284.

Pinetown: Isipingo North, Ward 354 (NU). Polela:

near Donnybrook, Reinhold in J 21535. Port

Shepstone: The Valleys Farm, Mogg 13947. Port

St. Johns: Brueckner & Allsopp 229 (NU).

Richmond: Byrne, Galpin 11989. Underberg:

Sani Pass, Hilliard s.n. (NU). Utrecht: Donkerhoek

Devenish 9.

Swaziland. — Mbabane: Forbes Reef Bush, Burtt

Davy 2744. Piggs Peak: Devils Bridge, Compton

31493.

Transvaal. — Barberton: Ameide Plantation,

Scheepers 1246. Belfast: Dullstroom, Galpin 13363.

Letaba: Grootbos Forestry Reserve, Scheepers 713.

Lydenburg: Sabie Falls, Legal in Col. Herb. 2455.

Petersburg: Blauwberg, Codd & Dyer 9095.

Pilgrims Rest: Graskop, Rogers 23029. Potgieters-

rust: Palala, Codd 2365. Soutpansberg: Entabeni,

Ohermeyer 1228. Wakkerstroom: Slangapiesberg,

Burtt Davy 1939. Waterberg: Geelhoutkop, Mogg

24500.

No justification was found for upholding var.

latior Pilg. as there are intermediates connecting it

with the typical form. All records of P. henkelii from

Swaziland and Transvaal have proved to be juvenile

specimens or coppice shoots of P. latifolius. For a

note on vegetative differences between the two

species see under P. henkelii.

The commonest member of the genus in Southern

Africa and one of the most abundant trees in the

Knysna forests. It yields valuable timber, and both

seeds and receptacles are eaten by birds and other

animals. Vernacular names used for the species are

True Yellowwood and Opregte Geelhout.

3. Podocarpus elongatus (Ait.) UHerit. ex

Pers., Syn. 2 : 580 (1807); Mirbel in Mem.

Mus. Hist. Nat. Paris 13 . 75 (1825); L.C. &

A. Richard, Comm. Bot. Conif. 13, t.l fig. 2

(1826); Endl., Syn. Conif. 218 (1847);

Carriere, Traite Conif. ed. 2 ; 671 (1867),

partly; Pari, in DC., Prodr. 16,2:511 (1868),

partly; Pilger in Pflanzenr. 4, 5 : 89 (1903),

not seen: Drege 6186, 6185, Bachmann 2211,

1522, 1523, Diels 937; Chalk et ah, Forest

Trees & Timbers Brit. Emp. 1 : 24 (1932);

Stapf in F.C. 5, 2 (Suppl.): 7 (1933), partly,

excl. Lucas s.n., Salisbury s.n. and Vere s.n.,

not seen: Bachmann 1522, 1523, 2211, Diels

937; Dallimore & Jackson, Handb. Conif.

ed. 3 : 66 (1948); Gray in Journ. Arn. Arb.

34 : 164 (1953), partly, excl. Zeyher s.n. in

part, not seen: Pearson 15551, “Hort.

Daudin”, Marloth 11488. Type: C.B.S.,

Masson s.n. (BM, lecto.!).

Taxus elongata Ait., Hort. Kew. ed. 1, 3 : 415

(1789); Thunberg, Prodr. 117 (1800); FI. Cap

ed. Schultes 547 (1823). Taxus capensis Lam.,

40

PODOCARPACEAE

Encycl. 3 : 229 (1789). Type: C.B.S., Sonnerat

s.n. in P 17135 (photo seen).

Podocarpus thunbergii var. angustifolia Sim, For.

FI. Cape Col. 332, t.149 fig. 3-5 (1907). Type:

the plate in Sim, l.c.

Rounded tree or spreading shrub usually

3-6 m high but attaining a height of 20 m;

diameter of shrubs as much as 12 m. Bark,

thin, more or less persistent, greyish-green to

dark grey. Branchlets pale yellowish-green,

terete with grooves from decurrent leaf bases.

Terminal buds of average branchlet about 2-3

mm in diameter; outer bud scales narrowly

triangular-oblong, 4-6 mm long and about

1-5 mm wide. Leaves spirally arranged to

subopposite, often crowded in upper parts of

shoots and subverticillate, spreading to sub-

erect, glaucous to greyish-green above, nar-

rowly oblong-elliptic, tapering in the upper

1/2 to 1/3, but more abruptly near the tip,

acute to subobtuse; adult leaves (1-8-)

3-6 (-7) cm long and (3-) 4-5 (-9)

mm wide; juvenile leaves up to 12 cm

long and 1 cm wide; midrib distinctly raised

on lower surface, on upper surface slightly

raised in lower half, margins flat to slightly

recurved; stomata on lower surface in

15-30 ± distinct longitudinal rows on either

side of midrib, on upper surface many leaves

have 1 to several short rows of stomata in

shallow longitudinal grooves. Male cones

solitary or in groups of 2-5, generally ± ses-

sile, more rarely on thin fertile shoots up to

7 mm long bearing 3-5 sessile cones in the

axils of bracts, cones (1-) 1-4-1 -9 (-2-5) cm

long, elongating up to 3 or 4 cm after shedding

pollen, (3-) 4 (-5) mm in diameter; outer

sterile scales at base trullate-ovate to broadly

transversely elliptic, contracted into a short

tip, margin brown, scarious, denticulate, 2-3

mm long and 1 • 5-2 mm wide; terminal lobe

of fertile scale triangular to ovate-triangular,

denticulate to lacerate, 0- 5-0- 6 mm long and

0-6-0 -8 mm wide; pollen sacs 0-9-1 -3 mm

long and 0-6-0 -7 mm in diameter. Female

cones solitary, on naked stalks (2-) 4—6 (-13)

mm long and about 1 mm in diameter;

receptacle fleshy, glaucous green at first, turn-

ing scarlet, when ripe 9-15 mm long and

10-16 mm wide, with 1 or 2 fertile scales, 1 or

2 seeds maturing on each receptacle. Seed

ellipsoid to ovoid, slightly apiculate, (6-) 7-10

(-12) mm long, dark glaucous green; total

thickness of shell 0-3-1 mm, consisting of 3

thin layers : the outer leathery, the middle one

slightly woody and the inner pergament-

aceous, mottled with dark purple, between

middle and outer layers small deposits of

yellow resin are common. Fig. 10 : 3.

Confined to the winter-rainfall region of the

western Cape where it grows mainly on sandy soil,

often along streams and rivers. It has not been

recorded from the coastal belt and appears to be

absent from the Cape Peninsula. In exposed positions

on mountains it is often stunted and can be almost

prostrate.

Cape. — Caledon: Kogelberg Reserve, Grobler in

STE 24247. Ceres: Thode A2284. Clanwilliam:

Cedarberg, F.D. Herb. 1243. Malmesbury:

Riebeeck Kasteel Rocks, Taylor 1568 (NBG).

Paarl: Paarl Mountain, Prior s.n. (K). Piquetberg:

Mouton Valley Farm, Marloth 11488. Robertson:

near Ashton, Marloth 11590. Stellenbosch: Swart-

boskloof, Kerfoot 5048. Swellendam: Bontebok

National Park, Barnard 666. Tulbagh: Witsenberg

Mountains, Marloth 1707. Van Rhynsdorp: Van

Rhyns Pass, Mauve 4148. Wellington: Berg River,

Marloth 11079. Worcester: Baineskloof, Compton

17950 (NBG).

May be confused with P. latifolius, but it is a

well-defined species and it is very unlikely that the

two species hybridise as Gray (Journ. Arn. Arb. 34 :

165, 1953) suggests. The presence of scattered stomata

on the upper surface of many leaves is a valuable

diagnostic character. The grooves in which these

stomata are situated are usually visible to the naked

eye and the stomata themselves can be seen under

10 x magnification. In the majority of specimens

20-80 per cent of the leaves have at least one short

row of stomata on the upper surface. Only a few

herbarium specimens were seen in which none of

the leaves had any stomata on the upper surface.

The two lectotypes chosen by Gray (l.c.) cannot

be accepted: firstly a species cannot be represented

by more than one collection and secondly there is

no indication that the author of the species saw

either of the specimens chosen. Therefore the Masson

specimen in BM has been chosen as the lectotype

(see Bothalia 9 : 142 (1966)).

Seeds and receptacles are eaten by birds and other

animals. The species is sometimes referred to as

Breede River Yellowwood.

4. Podocarpus henkelii Stapf ex Dallim.

& Jacks., Handb. Conif. 47 (1923), ( henckelii

sphalm.); Burtt Davy, FI. Transv. (1926),

partly, as to fig. 5 G only; Chalk et al., Forest

Trees & Timbers Brit. Emp. 1 : 23 (1932); l.c.

3 : 84 (1935); Stapf in F.C. 5, 2 (Suppl.): 9

(1933), partly, excl. Cooper 1928, Sim 19019,

Legat 2467 (3467 sphalm.), Nelson 420, Burtt

Davy 2748a (2738a sphalm.), not seen: F. D.

Herb. 1248, 1249, Henkel in F.D. Herb. 2331

a & b; Gray in Journ. Arn. Arb. 34 : 169

(1953), partly, excl. Legat 2467 ( Barbarton

2467 sphalm.), Burtt Davy 2748a, not seen:

Setchell s.n., Ad. No. 1282, “N.Y. Bot.

PODOCARPACEAE

41

Gard.”. Type: Mount Ayliff: Bulembu

Forest, Fort Donald Forest Station, Cochrane

in PRE 29648 (lecto.!).

Podocarpus thunbergii var. falcata Sim, Tree

Planting in Natal 236, 285, fig. 94 (1905), nom.

subnud.; sensu Sim, For. FI. Cape Col. 332, fig.

opp. p.55, 1. 1 49 fig. 1 (1907), partly, as to description

and figures but excl. synonymy. P.falcatus sensu

Sim, FI. Trees & Shrubs for Use in S. Afr. 185

(1919); Sim, Native Timbers of S. Afr. 102, fig. 1

(1921), non (Thunb.) R. Br. ex Mirb.

Straight tree up to 35 m high and attain-

ing a girth of up to 6 m. Bark dark grey to

khaki-coloured, in older specimens longi-

tudinally fissured, usually defoliating in long

narrow strips, occasionally defoliating in large

sheets. Branchlets pale green with grooves

from decurrent leaf bases. Terminal buds

3-5 mm in diameter; outer bud scales nar-

rowly triangular tapering to a i linear tip

0-5-1 mm long, 7—1 0 mm long and 1-4-1 -9

mm wide at the base, upper half often rolled

into a spiral. Leaves spirally arranged to sub-

opposite, often crowded in upper parts of

shoots and subverticillate, i pendulous, dark

green and glossy above, straight to slightly

falcate, very narrowly lanceolate-elliptic with

widest point generally just below the middle,

almost parallel-sided from there downward to

about the lower third, then narrowing gradu-

ally into a short petiole, tapering very

gradually from about the middle upwards to

an acuminate tip; adult leaves (5-) 9-12 (-17)

cm long and (5-) 6-5-8 (-10) mm wide;

juvenile leaves up to 22 cm long and up to 26

times as long as broad; midrib raised on

lower surface, on upper surface slightly raised,

at least in lower 2/3; margins slightly

reflexed; stomata confined to lower surface,

arranged in 25-35 fc distinct longitudinal

rows on either side of midrib. Male cones

solitary or in groups of 2-5, sessile or on very

short stalks, ( 1 - 2—) 1-5-1 -8 (-2) cm long,

elongating to 4 cm and more after shedding

pollen, (4-) 4 • 5-5 (-6) mm in diameter ; outer

sterile scales at base trullate-ovate to narrow-

ly so, often with a ± terete tip 0-25-0-75 mm

long, strongly but bluntly keeled, subentire to

finely lacerate, about 4-5 mm long and 2-3

mm wide ; terminal lobe of fertile scale widely

ovate to ovate-triangular, lacerate to irregu-

larly dentate, 0-7-1 mm long and 0-9-1 -1

mm wide, pollen sacs 1 -3-1-5 mm long and

0-6-0- 7 mm in diameter. Female cones

solitary on short naked stalks, receptacle

clavate, glaucous, never fleshy, often not

clearly demarcated from the stalk, length of

receptacle and stalk together 4-12 mm, width

of mature receptacle at top 3-6 mm, receptacle

consists of two bracts of which only the

terminal one is fertile. Seed ovoid-ellipsoid,

narrowing towards the base, ( 1 • 2—) 1-7-2 -2

(-2-5) cm long and (1 • 1-) 1 -4-1 -8 (-2- 1) cm

wide, olive-green; shell hard, leathery, gritty

but never woody, 1-4-4 mm thick, with

cavities up to 1 mm wide containing a pale

yellow to white resinous deposit. Fig. 10:4.

Occurs in montane and less commonly in coastal

forests in Natal and the adjoining regions of the

Eastern Cape.

Cape. — Engcobo: Engcobo Main Forest, Ryan

4469 (FFTO). Flagstaff: near Flagstaff. Hilliard s.n.

(NU). Mount Ayliff: near Mount Ayliff, Story 4248.

Mount Frere: near Mount Frere, Martin 108

(RUH). Tabankulu: Tonti Forest, Whibley in

F.D. Herb. 2167. Umzimkulu: Mhlonga Forest,

Kaufmann in F.D. Herb. 2170, 2171.

Natal. — Alfred: Impetyne Forest, Hilliard 1249

(NU). Bergville: Indumeni Forest, Killick & Marais

2144. Ixopo: Ingwangwane, Houshold in F.D. Herb.

1948. Pietermaritzburg: near Curry’s Post, Acocks

11578. Polela: near Donnybrook, Hilliard 2463

(NU). Port Shepstone: Glenmore, Van der Schijff

6646 (PRU).

As pointed out before, all records of P. henkelii

from Swaziland and Transvaal have proved to be

juvenile plants or coppice shoots of P. latifolius.

Sterile specimens of the two species can usually best

be distinguished as follows: in P. henkelii the widest

point of the leaf is usually about in the middle and

from here the leaf narrows gradually and evenly

to an acuminate tip; in P. latifolius leaves are

generally more or less parallel-sided in the middle

portion and taper fairly abruptly in the upper

1/4 or 1/5. Leaves of juvenile plants may be widest

in the middle but they too taper more or less abruptly

near the tip.

Tinley 734 and 745 (both in NU) were collected

in the Ngome forest in the Ngotshe district of

Natal, more than 100 miles from the nearest known

locality of the species. Their leaves show the charac-

teristic tapering but they are wider and shorter

than normal. Fertile material is needed for a definite

indentification.

A valuable timber tree often grown for orna-

mental purposes. It is commonly referred to as

Henkel’s Yellowwood.

42

PlNACEAE

PINACEAE

by J. P. Jessop

Trees or rarely shrubs, monoecious, evergreen or rarely deciduous, with monopodial

growth; the wood containing resin. Leaves linear, spirally arranged. Male strobiles small,

usually lateral, with numerous spirally arranged scales each bearing two pollen sacs on the

abaxial surface; pollen grains usually with a pair of wings; bracts absent. Female strobiles

usually subterminal, usually woody; scales arising in the axils of numerous spirally arranged

bracts and attached to them only at the base, bearing two inverted ovules on the adaxial

surface. Seeds usually winged; testa woody or leathery; aril absent. Cotyledons several to

numerous.

Nine genera recognised, containing about 200’ species, almost all occuring in the northern hemisphere.

Many species, mostly belonging to the genus Pinus, are cultivated in South Africa, two of which are natural-

ised and hence dealt with here. For a list of cultivated species see p.5\.

22 PINUS

Pinus L., Syst. Veg. 1000 (1753); Pilg. in Nat. Pflanzenfam. ed. 2, 13 : 165 (1926); Adamson

inFl. Cape Penins. 31 (1950); Loock, The Pines of Mexico and British Honduras (1950); Pilg.

& Melchior in Syllabus Pflanzenfam. 1 : 331 (1954); Dallimore & Jackson, Handbook of

Coniferae 442 (1961). Type species: P. sylvestris L.

Evergreen trees or shrubs often with apparently whorled branches. Branches of two

kinds; the main branches bearing only scale-like leaves lacking chlorophyll, and deciduous

short shoots bearing (1-) 2-5 (-8) green leaves in a common sheath. Green leaves needle-like,

usually with 2 or more resin canals. Male strobiles clustered at the base of the young twigs;

the pollen with bladder-like wings. Female strobiles terminal or lateral, ripening in their

second or third year; the scales becoming woody with the free ends expanded.

About 80, or more, species widespread in, and almost all restricted to, the northern hemisphere including

northern Africa. Two species are regarded as being naturalised in South Africa and are successfully com-

peting with the indigenous vegetation, viz: P. patula in the eastern Transvaal and P. pinaster in the southern

and south-western Cape Province. These two are fully described below. Many other species are cultivated

in South Africa (see /?.51).

Leaves arranged in bundles of 3, occasionally 4 or 5, pendulous 1 .P. patula

Leaves arranged in bundles of 2, patent 2. P. pinaster

1. Pinus patula Schlechtd. & Cham, in

Linnaea 6 : 354 (1831); Kotze, in S. Afr.

Journ. Sci. 23 : 453, plates III-V (1926);

Loock, The Pines of Mexico and British

Honduras (1950); Poynton, Notes on Exotic

Forest Trees in South Africa ed. 2 : 87 (1960) ;

Dallimore & Jackson, Handbook of Coni-

ferae 525 (1961).

Pyramidal trees up to about 40 m high.

Bark broken into longitudinal, irregular

shapes on the lower parts, but thin, papery

and reddish-brown higher. Young shoots with

many nodes, glabrous, glaucous-green at first

becoming brownish in the second year.

Winter buds cylindrical, long pointed, 12-25

mm long, covered with lance-shaped, long-

pointed, fringed scales. Leaves light green,

usually in bundles of 3 but sometimes 4 or 5,

pendulous, 15-22 cm long; basal sheath per-

sistent on the branches, about 25 mm long.

Female strobiles lateral, subsessile, in clusters

of 2-5, 7-10 cm long, ovoid-conic, slightly

curved, oblique at the base, pale brown, per-

sistent on the branches; scales smooth with

a minute prickle. Seeds triangular, grey

mottled with black, 5 mm long; wing 12-20

mm long.

Indigenous to east-central Mexico, this species

was introduced into South Africa in 1908. It now

covers the largest aiea of any coniferous species in

plantations, particularly in the eastern Transvaal,

PlNACEAE

43

where it is invading the vegetation on mountain

slopes, especially in the Sabie district. It is also

grown in the Natal Midlands and the eastern Cape.

O.F.S.— Harrismith: Harrismith Plantation,

Robertson in PRF 7848. t

Transvaal. — Ermelo: Jessievale Plantation,

without collector in PRF 5621.

This species is known as the Patula Pine. One

of its more characteristic features is the pendulous

nature of the leaves.

2. Pinus pinaster Ait., Hort. Kew. 3 : 367

(1789); Adamson in FI. Cape Penins. 32

(1950); Poynton, Notes on Exotic Forest Trees

in South Africa ed. 2 : 94 (1960); Dallimore

& Jackson, Handbook of Coniferae 528

(1961).

Pyramidal tree up to 40 m high. Bark

deeply fissured, reddish-brown. Young shoots

glabrous, pale brown, becoming prominently

ridged and roughened by the bases of scale

leaves. Winter buds stout, 20-25 mm or more

long, spindle-shaped, with whitish-brown,

fringed, reflexed scales. Leaves dark green, in

bundles of 2, rigid, 5-20 cm long. Female

strobiles subterminal, sessile, in clusters of

1 - many, 7-20 cm long, ovoid-conic, light

reddish-brown; scales keeled, with a promi-

nent, prickly umbo. Seeds 7-8 mm long with

a wing 25-35 mm long.

Indigenous to Mediterranean Europe, from

Portugal to Greece, and Africa as far south as

Algeria. It was probably introduced to the Cape in

the eighteenth century, and is now grown on a large

scale in the southern and south-western Cape. It

has become established in many parts of the

southern and south-western mountains and is

considered to constitute a serious threat to the

indigenous flora.

Cape. — Cape Town: Table Mountain, Marloth

s.n. (PRE). Knysna: Concordia Plantation,

Robertson in PRF 3001 . Wynberg: Tokai, Neethling

in PRF 6915.

This species is commonly known as the Cluster

Pine.

CUPRESSACEAE

by J. A. Marsh* *

Monoecious or occasionally dioecious, evergreen trees or shrubs; wood containing resin.

Leaves usually of two types, in juvenile form often needle-like, spirally or subspirally arranged

and spreading, in adult form usually small, squamiform, whorled or decussate and more or

less appressed. Male strobiles small, mostly terminal and solitary; scales few to many,

decussate or in whorls of three, bearing at the base on the abaxial surface 2-6 pollen sacs;

pollen grains not winged. Female strobiles developing into typical cones, with enlarged

leathery or woody valvate or imbricate and finally gaping scales, rarely fleshy and berry-like

( Juniperus , Arceuthos)-, scales few to many, all or most fertile, bearing erect ovules at the

base on the adaxial surface. Seeds free, rarely united in a stone, mostly winged; testa

crustaceous to woody. Cotyledons 2, rarely 3-6.

Phillips in his Genera of South African Flowering Plants ed. 2 (1951) placed IViddringtonia under Pinaceae

in the broad sense, but it has been decided to follow Pilger in Pflanzenfam. ed. 2, 13 : 361 (1926) and subse-

quent authors, such as Stapf in Flora Capensis 5, 2 (Suppl.) : 15 (1933), who take a more restricted view

of the family limits and include this genus in the family Cupressaceae.

Genera about 16, found in both hemispheres; 1 genus occurs in the area of the Flora. For cultivated

species see p. 52

t The abbreviation PRF has been accepted for the Forestry Department Herbarium, Pretoria, and should

replace "F.D. Herb.” used elsewhere in this publication.

* Submitted in partial fulfilment of the requirements for the degree of M.Sc. at the University of Pretoria.

44

ClJPRESSACEAE

38 WIDDRINGTONIA

Widdringtonia End/., Gen. PI. Suppl. 2 : 25 (1842); Syn. Conif. 31 (1847); Schlechtd. in

Linnaea 23 : 339 (1864-5); Parlatore in DC., Prodr. 16, 2 : 442 (1868); Mast, in Journ.

Linn. Soc. Bot. 37 : 207 (1905); Stapf in F.C. 5, 2 (Suppl.) : 15 (1933); Phill., Gen. ed. 2 : 51

(1951); Lewis in F.Z. 1, 1 : 86 (1960); Chapman in Kirkia 1 : 139 (1961).

Pachylepis Brongn. in Ann. Sc. Nat. ser. 1,30 : 190 (1833), non Lessing (1832); Endl., Gen. PI. 259 (1837).

Parolinia Endl., Gen. PI. Suppl. 1 : 1372 (1841), non Webb (1840).

Callitris sect. Pachylepis (Brongn.) Benth. in Benth. & Hook, f., Gen. PI. 3, 1 : 424 (1880).

Callitris sensu Eichler in Pflanzenfam. ed. 1, 2 : 94 (1889), partly; sensu Dur. & Schinz, Consp. FI. Afr.

5 : 951 (1894); Engl., Pflanzenw. Afr. 2 : 88 (1908).

Monoecious, evergreen trees or shrubs; wood fragrant, easily worked, containing oil

and resin. Leaves of two types, in juvenile form needle-like and subspirally arranged, in

adult form scale-like and appressed, decussate or alternate but not in whorls of 3 or 4. Male

cones small, c-4 mm long, terminal, mostly on short lateral branchlets; scales 5-7 pairs

(usually 6), decussate, coriaceous, peltate, with normally 4 pollen sacs at the base of each

scale. Female cones small, axillary, on elongated shoots, single or clustered; scales normally

4 of equal size and arranged in one whorl, rarely 5 or 6 scales, corky-coriaceous, apiculate,

divaricate at time of pollination, then closing; ovules several at the base of each scale; mature

cones woody, 1 -3-2-5 cm in diameter, opening at the apex, with 4 very thick, woody valves

often each with a prominent dor al cusp; exterior of valves smooth to warty. Seeds ovoid

or trigonous, winged; testa crustaceous. Cotyledons after germination 2, rarely 3, green,

needle-like.

Named after Captain Widdrington, R.N., F.R.S. (ne Samuel Edward Cook) who travelled in Spain from

1829-32 and published a book on European Pines (1843).

A small genus occurring mainly along the southern and eastern mountain ranges of Southern Africa and

extending as far north as Malawi (Nyasaland).

It is closely related to the North African genus Tetraclinis and the Australian Callitris; for differences

between them, see Chapman in Kirkia 1 : 138(1961). The genus Cupressus is superficially similar to Widdring-

tonia but differs from the latter in the fruit which consists of 6-8 peltate scales, each with a central boss or

triangular process on the outer surface.

Three species are recognised, all of which occur in Southern Africa.

Valves of female cone smooth to wrinkled, not tubercled along margin, if warty then irregularly so ; adult

leaves on ultimate branchlets narrowly oblong, subtriangular in transverse section

1 . W. cupressoides

Valves of female cone rough, with regular tubercles along margin ; adult leaves on ultimate branchlets

ovate, semi-circular in transverse section:

Seeds ovoid, triquetrous, obscurely winged; scar of attachment 4-5x6 mm; Clanwilliam district

2. W. cedarbergensis

Seeds somewhat flattened, conspicuously winged; scar of attachment 1 ■ 5 x 2-5 mm; Willowmore

district 3. W. schwarzii

1. Widdringtonia cupressoides ( L .) Endl.,

Cat. Hort. Vindob. 1 : 209 (1842); Syn. Conif.

33 (1847); Gordon, Pinet. ed. 1 : 333 (1858);

ed. 2 : 417 (1875): Parlatore in DC., Prodr.

16, 22 : 443 (1868); Mast, in J. Linn. Soc. Bot.

37 ; 270 (1905); Pilg. in Pflanzenfam. 13 : 383

(1926); Stapf in F.C. 5. 2 (Suppl.) : 18 (1933);

Adamson & Salter. FI. Cape Penins. 33

(1950); Chapman in Kirkia 1 : 138 (1961).

Type: No soecimer in Linn. Flerb.; the

original description is adequate.

Thuja cupressoides L., Mant. PI. 125 (1767);

Thunb., Prodr. 2 : 110 (1800); FI. Cap. ed. Schult.

500 (1823); Ait f., Hort. Kew. ed. 2, 5 : 322 (1813);

Harv., Gen. S. Afr. PI. ed. 1 : 31 1 (1838). T.

quadrangularis Vent. ex Duhamel, Trait,

des Arbr. 3:16 (1806). Type: “He de France au

Reduit”, Commerson in Herb, de Jussieu 17177

(P, holo. !).

Cupressus capensis Breon in Cat. PI. Cult. Jard.

Bot. et Nat. Bourb. ( 1 825). Type: “He de Bourbon”,

Breon s.n. (P, holo.!).

Pachylepis cupressoides (L.) Brongn. in Ann.

Sc. Nat. ser. 1, 30 : 190 (1833); Krauss, FI. Kap. u.

CUPRESSACEAE

45

Natal 1 : 152 (1846). P. commersonii Brongn. in

Ann. Sc. Nat. ser. 1, 30 : 190 (1833). Type: “lie

de France au Reduit”, Commerson s.n. in Herb, de

Jussieu 17177 (P, holo.!), isotype in Herb. Commer-

son (P!).

Callitris cupressoides Schrad. ex Drege, Zwei

Doc. 115, 126 (1843), nom. nud. C. cupressoides

(L.) Dur. & Schinz, Consp. FI. Afr. 5 : 950 (1894);

Marloth, Kapland 116, 198 ,199 (1908). C. natalensis

Endl. ex Fourcade in Rep. Natal For. 121 (1889),

nom. nud. C. commersonii (Brongn.) Dur. & Schinz,

Consp. FI. Afr. 5 : 950 (1894). C. whytei (Rendle)

Engl, in Pflzw. Afr. 2 : 89 (1908). C. mahoni

(Mast.) Engl, in Pflzw. Afr. 2 : 88 (1908).

Widdringtonia commersonii (Brongn.) Endl., Syn.

Conif. 34 (1847); Parlatore in DC., Prodr. 16, 2 : 443

(1868); W. natalensis Endl., Syn. Conif. 34 (1847);

Gordon, Pinet. 334 (1858); Parlatore in DC.,

Prodr. 16, 2 : 443 (1868); Dur. & Schinz, Consp.

FI. Afr. 5 : 951 (1894). Type: according to End-

licher, it is reputed to be a Krauss or Gueinzius

specimen from Port Natal. W. caffra Berg in

Bonplandia 8 : 190(1860). Type: no specimen cited.

W. whytei Rendle in Trans. Linn. Soc. Bot. 4 : 60

(1894); Mast, in Journ. Linn. Soc. Bot. 37 : 270

(1905); Stapf in F.T.A. 6, 2 : 334 (1917); Pilg. in

Pflanzenfam. ed. 2, 13 : 383 (1926); Stapf in F.C.

5, 2 (Suppl): 17 (1933); Lewis in F.Z. 1, 1 : 86

(I960); Chapman in Kirkia 1 : 138 (1961); White,

For. FI. N. Rhod. 8 (1962); Chapman, Veg.

Mlanje Mts., Nyas. (1962). Type: Mt. Mlanje,

Whyte s.n. (BM, holo.!). W. mahoni Mast, in

Journ. Linn. Soc. Bot. 37 : 271 (1905); Stapf

in F.T.A. 6, 2 : 334 (1917). Type: Melsetter,

Mahon ( K !). W. dracomontana Stapf in Kew Bull.

206 (1918); Chalk et al., Some E. Afr. Conif. &

Legum. 15 (1932); Stapf in F.C. 5, 2 (Suppl): 21

(1933); Chapman in Kirkia 1 : 138 (1961). Type:

Drakensberg Mts., Sanderson 2011 (K, holo.!; S!).

W. stipitata Stapf in Hook., Ic. Plant. 32 : 3126

(1930); F.C. 5, 2 (Suppl): 16 (1933); Chapman in

Kirkia 1 : 140 (1961). Type: Cult. Piet Retief,

Kotze 7048 (F.D. Herb. !).

Shrub or small tree up to 9 m high or, in

the tropics, attaining a height of about 50 m,

with stem diameter up to 2 m, crown pyramid-

al when young, spreading with age; bark

on young trees reddish grey, thin, fibrous,

flaking off annually. Leaves of two types;

juvenile leaves on seedlings and young trees

needle-like, spreading, 1-2 cm long and up

to 2 mm broad; adult leaves scale-like, ap-

pressed, narrowly oblong, often not strictly

decussate, subtriangular in transverse section,

adnate at the base, free and adnate portions

often equally long. Male cones mostly on

short lateral branchlets; scales usually 6

pairs, decussate, coriaceous, peltate, broadly

ovate with 4 pollen sacs at the base of each

scale. Mature female cones consisting of

usually 4 woody valves; valves smooth to

wrinkled, not tubercled along margin, if warty

then irregularly so. Seeds ovoid, conspicuous-

ly winged, somewhat flattened, outline in-

cluding wings oblong - obovate, wing retuse

at apex; seed scar about 1-5 x 2-5 mm.

Cotyledonary leaves (after germination) 20-25

mm long and about 2 mm broad. Fig. 11:2.

Found on mountain ranges of the south-eastern

and eastern Cape, Natal, Transvaal, extending into

Southern Rhodesia, Portuguese East Africa and

Malawi.

Cape. — Caledon: Hermanus, Smuts 1222.

George: Outeniqua Mts., Krauss 1140 (FI).

Grahamstown: Howiesonspoort, Zeyher 3885 (P);

Governorskop, Rogers 27668 (Z). Humansdorp:

Goukama, Burchell 5588 (K). Mt. Ayliff: Tonti,

Kriel 3004 (F.D. Herb.). Peninsula: Table Mt.,

Burchell 111 (K). Port Elizabeth: Otterford Reserve,

Rodin 1123 (K). Stellenbosch: Jonkershoek, Galpin

12789. Stockenstrom: Katberg, Muller s.n. Umtata:

Baziya, Sim 1473.

Natal. — Bergville: Cathedral Peak, Killick 1745.

Escourt: Cathkin Peak, Meebold 13144 (M);

Giants Castle, Sanderson 2011 (K).

Transvaal.- — Petersburg: Blaauwberg,

Schweickerdt 1821 (K); Wolkberg, Nanni s.n.

Pilgrims Rest: Mariepskop, Van der Schijff 4477.

Waterberg: Groothoek, Codd 3976. Soutpansberg:

Hanglip, Rodin 4050.

The Commerson specimens, reputedly from

Mauritius, on which W. commersonii was based,

were examined and found to be indistinguishable

from W. cupressoides. Mr. R. E. Vaughan of the

Sugar Industry Research Station, Mauritius, states

(personal communication) that no reference to

these specimens is to be found in Commerson's

manuscript notes and no species of Widdringtonia is

indigenous to the island. Therefore the Commerson

specimens were either obtained from a cultivated

plant (which is unlikely), or were collected at the

Cape and were accidentally incorporated into the

Commerson collection when it was shipped to

France after Commerson’s death.

W. stipitata is a form of W. cupressoides with

longer pedicels. As this characteristic is not constant,

this species is placed in synonymy.

Although an impression is gained that geo-

graphical races can be recognised in W. cupressoides

(sens, lat.), a range of variation is found in characters

such as leaf shape, seed and cone morphology.

Thus no reliable distinguishing character could be

found to separate the three “species” W. cupressoides,

W. dracomontana and W. whytei.

2. Widdringtonia cedarbergensis Marsh in

Bothalia 9 : 125 (1966). Type: Clanwilliam,

Cedarberg Mts., near Middelberg West Peak,

Luckhoff in PRE 29824 (PRE, holo.!).

Callitris arborea Schrad. ex Drege, Zwei Doc. 73

(1843), nom. nud.; Dallimore in Kew Bull.

1913 : 222 (1913); Stapf in F.C. 5, 2 (Suppl.): 24

(1933), in synon. C. ecklonii Schrad. ex Pappe, FI.

Cap. Med. Prodr. ed. 1 : 25 (1850), nom. subnud.

46

CUPRESSACEAE

Fig. 11. — 1, Widdringtonia cedarbergensis, branch with immature and old strobiles ( Walsh in PRE 29915);

la, anterior and posterior view of seed; lb, twig with male cones; lc, male cone, x 4; Id, juvenile

shoot (Gov. Forester in PRE 29914). 2, W. cupressoides, female strobile, smooth form ( Banks sub PRF

12818); 2a, female strobile, rough form (Forester, in PRE 29916); 2b, seeds showing different

forms. 3, W. schwarzii, seed (Gov. Forester in PRE 29917).

CUPRESSACEAE

47

C. juniperoides sensu Dur. & Schinz, Consp. FI.

Afr. 5 : 951 (1894); sensu Engl., Pflanzenw. Afr.

2 : 89 (1908); sensu Marloth, Kapland 167 (1908).

Widdringtonia wallichii Endl., Syn. Conif. 34

(1847), nom. nud.; Stapf in F.C. 5, 2 (Suppl.): 23

(1933), in synon. W. juniperoides sensu Endl.

Syn. Conif. 32 (1847); sensu Stapf in F.C. 5, 2

(Suppl.): 23 (1933); sensu C. S. Hubb. in S. Afr.

J. Sc. 33 : 572 (1937); sensu C. A. Smith in Journ.

S. Afr. For. Add. 25 (1955); sensu Chapman in

Kirkia 1 : 138 (1961). W. wallichiana Gordon,

Pinet., suppl. 107 (1875), nom. nud.

Tree normally 5-7 m high, occassionally

attaining a height of about 20 m with stem

diameter up to 2 m; crown pyramidal when

young, spreading with age; bark on young

trees reddish grey, thin, fibrous, flaking off

annually. Leaves of two types; juvenile leaves

on seedlings and young trees needle-like,

spreading, 1-2 cm long and up to 2 mm broad ;

adult leaves scale-like, appressed, ovate, 2-4

mm long, usually strictly decussate, semi-

circular in transverse section, adnate at the

base, free portion often much shorter than

adnate portion. Male cones 2 mm long, most-

ly on short lateral branchlets; scales usually

6 pairs, decussate, coriaceous, peltate, broadly

ovate, acuminate, with 4 pollen sacs at the

base of each scale. Mature female cones sub-

globose, about 2-5 cm in diameter, usually

consisting of 4 (rarely 5 or 6) woody valves;

valves rough, with regular tubercles along the

margin. Seeds ovoid, obscurely winged, tri-

gonous; seed scar about 4-5 x 6 mm.

Cotyledonary leaves (after germination) about

35 mm long and about 5 mm broad. Fig.

11:1.

Found on the Cedarberg Mts. near Clanwilliam

in the Cape Province; occurs singly or scattered

over some 30 miles on rocky outcrops at an altitude

of between 3,000 and 5,000 feet.

Cape. — Clanwilliam: Cedarberg Mts., Coetzee

s.n. ; Ecklon & Zeyher 74.3 (W); Forester s.n.

For reasons why it was found necessary to rename

and describe this species, see note under W. juni-

peroides (L.) Endl., nom. dubium (p. 48), and

Bothalia 9 : 124 (1966).

3. Widdringtonia schwarzii ( Marloth )

Mast, in Journ. Linn. Soc. Bot. 37 : 269

(1905); Marloth, FI. S. Afr. 1 : t,17D (1913);

Stapf in F.C. 5, 2 (Suppl.) : 22 (1933); Ward

in Journ. S. Afr. For. Ass. 31 (1958); Chap-

man in Kirkia 1 : 143 (1961); Liickhoff in

Bosbou S. Afr. 3 : 1 (1963). Type: Willow-

more, Marloth 3614 (PRE, holo.l; GRA;

NBG).

Callitris schwarzii Marloth in Bot. Jahrb. 36 : 206

(1905); Kapland 134 (1908).

Tree normally 17-26 m high, occasion-

ally attaining a height of about 40 m with stem

diameter up to 5 m; crown pyramidal,

usually not spreading with age; bark on

young trees reddish grey, thin, fibrous, flaking

off annually. Leaves of two types; juvenile

leaves on seedlings and young trees needle-

like, spreading, 1-2 cm long and up to 2 mm

broad; adult leaves scale-like, appressed,

ovate, usually strictly decussate, semi-circular

in transverse section, adnate at the base, free

portion often much shorter than adnate por-

tion. Male cones mostly on short lateral

branchlets, scales usually 6 pairs, decussate,

coriaceous, peltate, oblong, acuminate, with

4 pollen sacs at the base of each scale. Mature

female cones consisting of usually 4 woody

valves, valves rough with regular tubercles

along the margin. Seeds ovoid, conspicuously

winged, somewhat flattened, outline including

wings oblong-obovate, wing retuse at apex;

seed scar about 1-5 x 2-5 mm ; Cotyledonary

leaves (after germination) 20-25 mm long and

about 2 mm broad. Fig. 11:3.

Found in rocky ravines of the Baviaanskloof and

Kouga Mts., Willowmore district, Cape Province.

Cape. — Willowmore: Civil Commissioner s.n.

(K); Baviaanskloof Mts., Forest Officer in F.D.

Herb. 7157 (F.D. Herb.); Kouga Mts., Burton

4019 (F.D. Herb.); Gov. Forester 2; 4 (UPR);

Sim 2920 (BOL).

Doubtful and Excluded Species

Cupressus juniperoides L., Sp. PI. ed. 2 : 1422

(1763), was based on young seedlings grown

from seed said to have come from “Cap. b.

Spei.” From the description it is impossible

to identify the species or even the genus.

There is no record that any subsequent author

saw a specimen and no specimen is present in

the Linnaean Herbarium. Judging from speci-

mens seen in the British Museum and Geneva,

the name was applied by botanists before 1800

to the S.W. Cape plant described by Linnaeus

as Thuja cupressoides in 1 767 . At the time when

Linnaeus described Cupressus juniperoides,

the Cedarberg Mts. were unexplored

botanically. In view of the above evidence, it

is considered advisable to reject the name as

a nomen dubium.

Cupressus africana Mill., Gard. Diet. ed. 8

(1768), nom. dubium, is presumbly W. cupres-

soides from the Cape Peninsula.

Juniperus capensis Lam., Encycl. Meth. 2 : 626

(1786), nom. dubium. The type, which was

examined in the Herbarium of the Museum

48

CUPRESSACEAE

National d’Histoire Natureile, Paris, consists

of sterile, juvenile branches, which could not

be identified with any degree of certainty.

Schubertia capensis Spreng., Syst. Veg. 3 : 890

(1826), nom. illegit., based on Cupressus

juniperoides L. (q.v.).

Pachylepis juniperoides (L.) Brongn. in Ann. Sc.

Nat. 30 : 190 (1833), nom. dubium, based on

Cupressus juniperoides L. (q.v.).

Widdringtonia juniperoides (L.) Endl., Syn.

Conif. 32 (1847), nom. dubium, based on

Cupressus juniperoides L. (q.v.).

Parolinia juniperoides (L.) Endl. ex Gordon,

Pinet. ed. 2 : 418 (1875), nom. dubium, based on

Cupressus juniperoides L. (q.v.).

Callitris juniperoides (L.) Dur. & Schinz, Consp.

FI. Afr. 5 : 951 (1894), nom. dubium, is also

based on Cupressus juniperoides L. (q.v.).'

Widdringtonia equisetiformis Mast, in J. Linn.

Soc. Bot. 37 : 271 (1905) was based on a culti-

vated specimen from Tokai in the Cape. It

was later placed as a synonym of the Australian

species Callitris robusta R. Br. by Masters

himself in J. Linn. Soc. Bot. 37 : 332 (1905).

Nomina Nuda

The following nomina nuda or subnuda cannot

be assigned with certainty to any recognised species.

Callitris strict a Schlechtd. ex Hook, in Lond.

J. Bot. 4 : 141 (1845).

Widdringtonia ericoides Knight in Syn. Conif.

PI. 13 (1850).

Callitris capensis Schrad. ex Gordon, Pinet.

333 (1858).

Widdringtonia glauca Carr., Trait. Conif. ed.

2 : 61 (1867).

WELWITSCHIACEAE

by I. C. Verdoorn

Plant dioecious, stem woody, unbranched (closed system of growth), broadly obconic

or turbinate, concave on top, outer covering corky; taproot elongated, branching near the

apex. Leaves produced from the subcircular groove around the rim of the crater-like apex

of the plant, parallel-veined, persistent, growing from the base. Inflorescences arising from

floriferous cushions above, sometimes also below, the insertion of the leaf. Flowers unisexual

(the male pseudo-bisexual but functionally unisexual), borne singly in the axils of scales

forming small cones. Male flower: perianth segments 4, in 2 whorls; stamens 6 with filaments

connate at the base; anthers 3-celled dehiscing by 3 slits at the summit; ovule single, abortive,

with the integument produced into a style-like, tubular structure and expanded at the apex

into a stigma-like, papillose disc. Female flower: perianth segments 2, navicular, united along

the margins to enclose the ovule, flattened, with wings developing along the keels;

ovule with the single integument produced into a style-like tubular structure which eventually

is exserted, widened and lobed at the apex. Seeds winged, with 2 cotyledons, endospermous.

Family with 1 genus and 1 species, found in Africa.

The single genus in this family was at first placed in the Gnetaceae, the family under which it appears in

Phillips’s Genera of South African Flowering Plants, ed. 2 (1951). Because of the striking morphological

and physiological differences between Welwitschia and related genera, as well as the unique form of the

plant, it has been decided to follow Markgraf who, in Engler & Prantl’s Pflanzenfamilien ed. 2, 13 : 407

(1926), gives each of the genera, formerly in Gnetaceae, the status of family. This concept is today generally

accepted, for example by Pearson in the Flora Capensis, Vol. 5, 2 (Suppl.) : 1 (1933) and Lawrence, Taxonomy

of Vascular Plants (1960).

Among the unique characteristics found in Welwitschia are the following: — a closed system of growth;

a single pair of leaves which persist for the life-time of the plant; male flowers with a rudimentary ovule,

the integument of which is produced into a style-like structure which expands at the apex into a stigma-like

disc; and winged seeds.

Welwitschiaceae

49

48 WELWITSCHIA

Welwitschia Hook.f. in Gard. Chron., 71 (1862); in Trans. Linn. Soc. 24 : 6, tt.1-14 (1863);

Marloth, FI. S. Afr. 1 : 107 (1913); Pearson in F.C. 5, 2 (Suppl.) : 1 (1933); Phill., Gen. ed.

2 : 52 (1951); nom. cons. Type species: Welwitschia bainesii (Hook, f.) Carr.

Tumboa Welw. in Gard. Chron., 75 (1861); in Journ. Linn. Soc. Bot. 5 : 185 (1861). Type species: Tumboa

strobilifera Welw. ex Hook.f.

Description as for family.

Only one species is known, limited in distribution to desert or desert margin along the west coast of

Africa from the Kuiseb river, which runs into Walvis Bay, South West Africa, northwards to and slightly

beyond. Cape Negro in Angola.

The genus was first described in 1861 as Tumboa by Welwitsch, who had discovered these strange plants

near Cape Negro in Angola. The name Tumboa was derived from what Dr. Welwitsch had been told was

the native name for these plants. A year later, with Welwitsch’s consent, J. D. Hooker changed the generic

name to Welwitschia. According to present day rules of nomenclature this required official conservation

and in the I.C.B.N. (1935), on page 89, Welwitschia Hook.f. appeared in the list of conserved names.

Welwitschia bainesii {Hook.f.) Carr.

Conif. ed. 2 : 783 (1867); Pearson in F.C. 5,

2 (Suppl.) : 1 (1933); Whellan in Kirkia 5,

1 : 33 (1965). Type: Damaraland, Baines s.n.

(K, photo, PRE).

Tumboa bainesii Hook. f. in Gard. Chron.

1007 (1861); Naud. in Rev. Hort. 186 (1862) as

Toumboa; Rendle in Cat. Afr. PI. Welw. 2 : 257

(1899). Type as above. T. strobilifera Welw. ex

Hook.f. in Gard. Chron. 71 (1862). Type: Angola,

Cabo Negro, Welwitsch s.n. (K, photo, PRE).

Welwitschia mirabilis Hook.f. in Gard. Chron. 71

(1862); in Trans. Linn. Soc. Bot. 24 : 7, tt.1-14

(1863); in Bot. Mag. tt.5368, 5369 (1863); McNab

in Trans. Linn. Soc. Bot. 28 : 507, t.40 (1873);

Sykes in Trans. Linn. Soc. Bot. ser. 2, 7 : 327

(1910); Pearson & Prain in F.T.A. 6, 2 : 333 (1917).

Type: Angola, Cabo Negro, Welwitsch s.n. (K,

photo, PRE).

Plant body unbranched, woody, obconic

or turbinate, about 60 cm long, rarely up to

2 m long, usually with only the crater-like

apical portion above ground, diam. at apex

0-3-1 -2 m, outer covering on exposed part

corky, corrugated; taproot elongated,

0-3-1 m long, sometimes up to 3 m long,

usually branching only towards the apex.

Leaves 2, rarely 3, produced from a groove

encircling the rim of the crater-like apical

portion, oblong, broadly acuminate to a sub-

acute apex, 2-3 m long, at base as broad as

about half the circumference of the plant apex,

parallel-veined, with unusual lateral vascular

communication, persistent, growing from the

base, with age tearing into strips. Inflorescence

a complex cyme, several to many arising

annually from pits in cushions along the leaf

bases, 1 5-30 cm long, bearing several to many

cones; male cones deep salmon coloured

when mature with many imbricate scales,

stamens eventually exserted, subquadrate,

1-4 cm long, 5-7 mm broad; female cone

glaucous green becoming yellowish and

banded with salmon red at maturity, scales

tightly imbricating, style-like structure eventu-

ally exserted, subquadrate, 2-8 cm long,

1 - 5—3 cm broad. Flowers unisexual and

dioecious, borne singly in the axil of the

scales; male: segments 4, 2 outer navicular,

about 4 mm long, 2 inner broadly obovate,

4 mm long, 5 mm broad near apex, filaments

united into a tube 3 mm long, free for about

1 • 5 mm, anthers terminal, oblate, 3-celled,

dehiscing by 3 terminal slits; ovule with the

integument produced into a style-like tube

with a stigma-like disc at the apex; female:

segments 2, naviculate and fused along their

margins to enclose the ovule, eventually

winged along the keels, persistent, ovule with

the integument produced into a style-like,

slender tube, fimbriate at the mouth. Seed

dicotyledonous, endospermous, enclosed in

the persistent membranous winged perianth

segments. Chromosomes: 2 n = 42 (Chromo-

soma 14 : 522-533, 1963). Fig. 12.

Found mainly in the desert or semidesert areas

along the west coast of South West Africa and

Angola, which is known as the northern section

of the Namib Desert. These areas stretch from the

Gobabeb Research Station on the Kuiseb River in

the south to just north of Cape Negro in Angola,

that is a few miles south of Mossamedes. Favours

dry watercourses in the desert localities but also

found on rocky ledges in the Swakop gorges.

Inhabits coarse gravelly soils (not deep sand as

generally believed). In localities farther inland such

as near the Petrified Forest nearly one hundred

miles from the coast, associated with Colopliosper-

mum mopane in, usually, loamy soils.

S.W.A. — Swakopmund: Dinter 6792; 35 miles

east of Swakopmund, Hardy & de Winter 1428

a and b, 2016 a and b; Heikamkab, lower Swakop

valley, Marloth 1191; upper portion of ravines

50

Welwitschiaceae

Fig. 12. — 1, Welwitschia bainesii, male plant; a, portion of male inflorescence with two male cones; b, male

flower with one of the inner segments pulled down to show the filaments united into a tube, x 5; c,

anther, x 10; d, integument enclosing the ovule and produced into a style-like tube, X 7 ( Hardy & De

Winter 2016a); e, portion of female inflorescence with two cones; f, female flower, nearly mature,

showing the two segments winged along the keels and the very slender style-like tube of the integument

exserted at the apex, x ( Hardy & De Winter 2016b).

Welwitschiaceae

51

leading into main valley, Galpin & Pearson 7440;

7470; 7476; Walvis Bay, Cleverly 1725; Kock s.n.;

east of Goanikontes, Rodin 2606. Omaruru: Brand-

berg, Strey 2212a; 2212b; 2212c. Outjo: Giess

3228; 3857; 3867. Kaokoveld: Otjikongo,

Merxmiiller 7447.

The largest known specimens of Welwitschia are

found in the desert areas such as the Welwitschia

Flats near Swakopmund. Nearer the escarpment,

where the rainfall increases, the average size of the

plants decreases but they are usually much more

abundant here than in the true desert.

CULTIVATED GYMNOSPERMS

by R. J. Poynton

Most Gymnosperms of economic value in other parts of the world have been introduced

into South Africa. Many have grown very fast and now form the basis of the softwood saw-

timber industry in this country; others are widely planted for shelter and ornament.

Representative examples are given below. All voucher specimens cited are housed in the

Herbarium of the Forest Research Institute, Pretoria (PRF).

GINKGOACEAE

GINKGO L.

Ginkgo biloba L., Maidenhair Tree, is widely,

though sparingly cultivated for ornament. Trans-

vaal.— Pretoria: Zoological Gardens, Poynton in

PRF 13680.

ARAUCARIACEAE

AGATHIS Salisb.

Agathis brownii (Lem.) L. H. Bailey [A. robusta

(C. Moore) F. Muell.] has been grown on a very

small scale for timber in Zululand and is of potential

economic importance. It is also sometimes en-

countered as an ornamental tree. Natal. —

Mtunzini: Port Durnford, Van Rensburg in PRF

10999.

Agathis microstachya J. F. Bailey & C. T. White

has been planted experimentally for timber in

Zululand and may be grown for this purpose on

a larger scale in the future. Natal. — Lower

Umfolozi: Kwambonambi, Haigh in PRF 13033.

Agathis palmerstonii F. Muell. has been planted

for timber on an experimental scale in Zululand

and may be grown more extensively in the future.

Natal. — Lower Umfolozi: Kwambonambi, Haigh

in PRF 13032; Sherry in PRF 13362.

ARAUCARIA Juss.

Araucaria angustifolia ( Bertol .) Kuntze (A. bra-

si liana A. Rich.), Parana Pine, has been sparingly

planted for timber and ornament in warm, humid

parts of the summer rainfall area. Cape. —

Mquanduli: Wilo, without collector, in PRF 11201.

Araucaria bidwillii Hook., Bunya-Bunya, is grown

as an ornamental tree in warm areas. Cape.—

Engcobo, without collector, in PRF 3742.

Araucaria columnaris (Foist.) Hook. (A. cookii

R. Br.), Captain Cook’s Pine, is commonly grown

for ornament, especially along the Natal coast.

Cape. — Uitenhage: Magennis Park, without col-

lector, in PRF 4212, 4237—8.

Araucaria cunninghamii D. Don, Moreton Bay

Pine, has been grown experimentally for timber in

warm, humid parts of the summer rainfall area

and is of potential economic importance in Zululand.

Transvaal. — Pretoria: Kotze in PRF 4029.

Araucaria heterophylla (Sulisb.) Franco [A. excelsa

(Lamb.) R. Br.], Norfolk Island Pine. Sparingly

grown as an ornamental tree in all warm areas.

Cape. — George: Simmonds in PRF 4005.

PINACEAE

PINUS L.

Pinus ayacahuite Ehrenberg, Mexican White Pine,

together with var. brachyptera Shaw, is of potential

value as a timber tree in the colder parts of the

summer rainfall area. O.F.S. — Harrismith: Kotze

in PRF 5106.

Pinus canariensis C. Sm., Canary Pine, is grown

extensively in the Cape Province for the production

of poles and is also planted for shelter, shade and

ornament. Transvaal. — Pretoria: Loock in PRF

9992.

52

Cultivated Gymnosperms

Pinus caribaea Morelet, Caribbean Pine, is grown

for timber in Zulnland and may be planted on a

much larger scale in the future. Natal. — Mtunzini:

Port Durnford, Liickhoff in PRF 10267.

Pinus elliottii Engelm ., Slash Pine, is grown on

a very large scale in moist parts of the constant and

summer rainfall areas for timber. In Zululand it is

spreading naturally to some extent. Transvaal. —

Pilgrim’s Rest: Tweefontein, without collector, in

PRF 12015.

Pinus khasya Royle ( P . insularis Endl.) has been

planted experimentally for timber and is of potential

economic importance in warm, humid parts of the

summer rainfall area. Transvaal. — Louis Trichardt :

Hanglip, Robertson in PRF 7850.

Pinus montezumae Lamb, has been planted for

timber on a small scale in cool, moist parts of the

summer rainfall area. Transvaal. — Belfast:

Robertson in PRF 7871.

Pinus palustris Mill., Long-leaf Pine, has been

planted fairly extensively for timber in moist parts

of the constant and summer rainfall areas. Trans-

vaal.— Nelspruit: Coetzeestroom, Liickhoff'm PRF

13588.

Pinus patula and P. pinaster (see pp. 42-43).

Pinus pinea L., Stone Pine, is a common orna-

mental tree in the western Cape Province, where

it propagates itself to a certain extent. Cape. —

Cape Town: Walsh in PRF 13681.

Pinus pseudostrobus Lindl. has been grown for

timber on a small scale under cool, moist conditions

in the summer rainfall area and is likely to be more

extensively planted in the future. Transvaal. —

Louis Trichardt: Hanglip, Robertson in PRF 7825.

Pinus radiata D. Don (P. insignis Dough),

Monterey Pine, is planted for timber and shelter

in the Cape Province. Cape. — Bredasdorp : Napier,

Scheuble in PRF 12527.

Pinus roxburghii Sarg. ( P . longifolia Roxb. ex

Lamb.), Chir Pine, has been grown extensively for

timber in the summer rainfall area and is still

planted for shelter and ornament, expecially in dry

localities. Transvaal. — Pretoria, Keet in PRF 8304.

Pinus taeda L., Loblolly Pine, is grown fairly

extensively for timber in cool, moist parts of the

constant and summer rainfall areas. Transvaal. —

Ermelo: Jessievale, without collector, in PRF 5476.

CEDRUS Link

Cedrus deodara (Roxb. ex Lamb.) G. Don, the

Deodar, is grown on a small scale for timber in

cool, moist parts of the summer rainfall area and

is widely planted for shelter and ornament. Trans-

vaal.— Pretoria: Bryntirin, Legat in PRF 4987.

PICEA A. Dietr.

Picea smithiana (Wall.) Boiss., West Himalayan

Spruce, has been planted experimentally for timber

in cool, moist parts of the summer rainfall area and

is sometimes grown for ornament. Natal. — Lions

River: Karkloof, Poynton in PRF 12838.

ABIES Mill.

Abies religiosa Schlechtd. <& Cliam., Sacred Lir, has

been planted experimentally for timber in cool,

moist parts of the summer rainfall area. Cape. —

Victoria East: Hogsback, without collector, in PRF

9744.

TAXODIACEAE

CRYPTOMERIA D. Don

Cryptomeria japonica (L.f.) D. Don, Japanese

Cedar, is grown on a limited scale for timber in

cool, moist parts of the summer rainfall area and

is sometimes encountered as an ornamental tree.

Transvaal. — Louis Trichardt: Hanglip, Holtz-

kampf in PRF 5449.

TAXODIUM Rich.

Taxodium distichum (L.) Rich., Swamp Cypress,

is widely, though sparingly cultivated for ornament.

Cape. — Cape Town: Botanic Gardens, Legat in

PRF 4033.

CUPRESSACEAE

CALLITRIS Vent.

Callitris arenosa A. Cunn. ex Mirb. (A. columel-

laris F. Meull.) is of potential value as a timber

tree, especially in Zululand. Transvaal. — Louis

Trichardt: Hanglip, Poyton in PRF 11965.

Callitris calcarata (A. Cunn.) R. Br. [ C . endlicheri

(Pari.) F. M. Bailey], Black Callitris, is sparingly

cultivated for shelter and ornament. Transvaal. —

Letaba: Woodbush, Kotze in PRF 2508.

Callitris glauca (R. Br. ex Mirb.) R. T. Bak. &

H. G. Sm. [ C . hugelii (Carr.) Franco], White

Callitris, has been grown on a very small scale for

timber in moist, warm parts of the summer rainfall

area and is sparingly cultivated for shelter. Trans-

vaal.— Lichtenburg: Poynton in PRF 12002.

Callitris rhomboidea R. Br. ex A. & L. C. Rich.

[C. cupressiformis G. Don: C. tasmanica (Benth.)

R. T. Bak. & H. G. Sm.] has been planted experi-

mentally for timber in warm areas and is occasionally

grown for ornament. Natal. — Mtunzini: Port

Durnford, Haigh in PRF 13021.

THUJA L.

Thuja orientalis L., Chinese Arbor-vitae, is

widely, but sparingly grown for ornament and as

a hedge plant. Transvaal. — Letaba: Grenshoek,

Bridgeford in PRF 10709.

Cultivated Gymnosperms

53

CUPRESSUS L.

Cupressus funebris Endl., Weeping Cypress, is

occasionally planted for ornament. Transvaal. —

Pretoria: Poynton in PRF 12695.

Cupressus glabra Sudw. has been much planted

for shelter and as a hedge plant. It hybridises

freely with C. arizonica Greene and is frequently

mistaken for it. Transvaal. — Belfast: Poynton

in PRF 11397-8.

Cupressus lusitanica Mill. (C. lindleyi Klotzsch),

Mexican Cypress, has been planted on a small

scale for timber in cool, moist parts of the summer

rainfall area and may be grown more extensively

in the future. Transvaal. — Ermelo: Jessievale,

Poynton in PRF 11407-09.

Cupressus macrocarpa Hartweg, Monterey

Cypress, is grown for shelter and ornament, more

especially in the western Cape. Cape. — Wynberg:

Tokai, Poynton in PRF 12698.

Cupressus sempervirensL., Mediterranean Cypress,

is widely cultivated for ornament — more especially

the fastigiate form, var. stricta Ait., the Churchyard

Cypress. Cape. — Wynberg: Tokai, Poynton in

PRF 12591.

Cupressus torulosa D. Don, Himalayan Cypress,

has been planted experimentally for timber in moist

parts of the summer rainfall area. Cape. — -Wynberg:

Claremont, Legat in PRF 4239.

JUNIPERUS L.

Juniperus virginiana L., Pencil Cedar, has been

planted experimentally for timber under a very

wide range of conditions and is occasionally en-

countered as an ornamental shrub or tree. Cape. —

Robertson: Everest in PRF 1090-2.

TYPHACEAE

by J. G. Anderson

Monoecious perennial herbs with creeping rhizomes. Stems erect, simple, terminated by

dense cylindrical flower-spikes. Leaves mostly radical, distichous, elongate-linear. Inflorescence

a dense spike of closely packed flowers with the male flowers zoned in the upper portion

contiguous to or shortly separated from the female flowers in the lower portion. Flowers

minute. Male flowers usually subtended by variously shaped scales or bracteoles; perianth

ab ent or of 3-6 small scales; stamens 2-5; filaments free or variously connate; anthers

linear, basifixed. Female flowers ebracteolate or with slender clavate or spathulate bracteoles,

abortive clavate female flowers (carpodia) often produced; perianth of several fine, persistent,

filiform or clavate hairs; ovary superior, often stalked, fusiform, unilocular with a solitary

pendulous ovule; style elongated, slender; stigma linear or lanceolate. Fruits minute,

ellipsoid or subcylindrical. Seed with a striate testa and mealy endosperm; embryo narrow,

nearly as long as the seed.

According to Hutchinson, Fam. Flow. PL 2 : 637 (1959), the Typhaceae is a monogeneric family, and

this view is accepted here.

49 TYPHA

TyphaE., Sp. PI. 971 (1753); Gen. PI. ed. 5:418 (1754); Rohrb. in Verh. Bot. Verein. Brandenb.

11 : 67 (1869); Benth. & Hook, f., Gen. PI. 3 : 954 (1883); Kronf. in Verh. Zool-Bot. Ges.

Wien 39 : 89 (1889); Engl, in Pflanzenfam. 2, 1 : 183 (1889); N.E. Br. in F.C. 7 : 31 (1897);

Graebn. in Pflanzenr. 4 : 8 (1900); N.E. Br. in F.T.A. 8 : 134 (1902); Phill., Gen. ed. 2 : 52

(1951); Hutch., Fam. Flow. PI. 2 : 637 (1959). Type species: T. angustifolia L.

Description as for the family.

Aquatic or marsh plants, widely distributed throughout the world, and comprising approximately 25

species, some of which are probably only forms of widespread variable species. The genus is represented

by only one subspecies in South Africa.

The name Typha is derived from the Greek word tuphos, a marsh.

54

Typhaceae

Typha latifolia L. subsp. capensis Rohrb.

in Verh. Bot. Yerein. Brandenb. 11:96 (1869) ;

Graebn. in Pflanzenr. 4, 8 : 10 (1900), nom.

superfl. Type: Cape, Uitenhage, Zwartkops-

river, Ecklon & Zeyher 913 (SAM, lecto.!).

T. latifolia L. forma capensis Rohrb. ex Kronf. in

Verh. Zool-Bot. Ges. Wien 39 : 180 (1889), in

syn. T. capensis (Rohrb.) N. E. Br. in F.C. 7 : 32

(1897). T. australis sensu N. E. Br. in F.C. 7 : 32

(1 897) excl. syn., non Schum. & Thonn. T. angusti-

folia L. p australis Rohrb. in Verh. Bot. Verein.

Brandenb. 11 : 83 (1869), partly as to S. Afr. speci-

mens. T. angustifolia L. subsp. australis Graebn. in

Pflanzenr. 4:13 (1900), partly as to S. Afr. specimens.

Perennial, robust, up to 2 m or more tall,

glabrous. Stems erect, simple. Leaves dis-

tichous but spirally twisted at the base and

sheathing forming a pseudo-stem, linear,

tapering to an obtuse apex, convex on the

back and flat or slightly concave below on

the inner surface, thinner and flat towards

the apex, 40-150 cm long and 4—20 mm broad,

margins smooth, often undulated in the upper

half, green or glaucous. Inflorescence a dense

cylindrical brown or yellow-brown spike

terminating the stem; upper male portion

15-30 cm long; lower female portion shorter,

contiguous with the male or up to 4 cm dis-

tant. Male flowers usually with a few various-

ly shaped, linear, lanceolate, cuneate or

forked, brownish-white, slender bracteoles;

stamens with slender filaments bearing 2 or 3

anthers; anthers 4-5 mm long, linear, often

distinctly twisted, 2-celled, pollen simple or

rarely compound. Female flowers usually

ebracteolate occasionally with a few slender,

colourless or brownish, lanceolate, clavate

or linear, acute or acuminate bracteoles,

abortive female flowers (carpodia) often pro-

duced; perianth of several, slender, colour-

less, simple hairs; ovary superior, usually

stalked, often brown-mottled, narrow, unilo-

cular with a solitary pendulous ovule; style

slender; stigma lanceolate or linear, brown.

Fruits minute, ellipsoid or subcylindrical,

about 1 mm long and 0-5 mm wide at the

middle. Fig. 13.

Widely distributed in Southern Africa and also

recorded from Angola, Rhodesia and Mozambique.

It occurs along stream-banks and in marshy places.

Cape. — Barkly West: Newlands, Wilman s.n.

Bathurst: Fish River, at mouth of Kap River,

Dyer 3388. Humansdorp: Fourcade 1778. Komga:

Flanagan 979. Middelburg: Grootfontein, Theron

1676. Peninsula: Diep River, Marloth 7290; 7291;

Observatory, Marloth 7289; Cape Flats, Andreae

243. Port Elizabeth: mouth of Swartkops River,

Theron 1648. Queenstown: Borigola, Galpin 8416.

Riversdale: Oakdale, Muir 3060. Uitenhage: Near

Prentice Kraal, Bolus 1469. Worcester: Veld

Reserve, Van Breda 60.

Natal. — Durban: Nelson 11745. Hlabisa:

Hluhluwe Game Reserve, Ward 2104. Inanda:

near Umhlanga Rocks Hotel, Wat mo ugh 489.

Umlazi: Isipingo Beach, Ward 619.

O.F.S. — Kroonstad: Pont 691. Thaba Nchu:

Maroka School, Potts 1929.

Basutoland. — Feribe: Dieterlen 18. Maseru:

Guillarmod 2906.

Swaziland. — Hlatikulu: Kubuta, Compton

28123.

Transvaal. — Nelspruit: Codd 8192. Pretoria:

Leendertz 687. Waterberg: Naboomspruit,

Mosdene, Galpin M.373.

S.W.A. — Grootfontein: Gautscha Pan, Story

5302.

Rohrbach in Verh. Bot. Verein. Brandenb.

1 1 : 96 (1869) states that he regards the South

African specimens, with which he was dealing, as

constituting a subspecies of T. latifolia F. He,

however, refers to the group as “T. capensis”,

that is, in binomial form. According to Art. 24 of

the International Rules of Botanical Nomenclature,

1961, this must today be rendered as T. latifolia F.

subsp. capensis Rohrb. Certain authors have

inferred from Rohrbach's publication that he esta-

blished T. capensis as a species. This led to N. E.

Brown unintentionally but validly effecting a change

of status for this name when in the Flora Capensis

he gave it specific rank for which the correct citation

is T. capensis (Rohrb.) N. E. Br. From the above

it follows too that the combination T. latifolia L.

subsp. capensis (Rohrb.) Graebn. in Pflanzenreich,

Typhaceae: 10 (1900) is superfluous. T. latifolia F.

forma capensis Rohrb. is published by Kronfeld

in Verh. Zool-Bot. Ges. Wien 39 : 180 (1889)

only as a synonym of T. latifolia L. subsp. capensis.

After examining all available South African

herbarium material of Typha, as well as fresh

material, it was concluded that only one taxonomic

unit occurs in South Africa. N. E. Brown in Flora

Capensis 7 : 31 (1897) included two species, T.

australis and T. capensis, separated on the presence

or absence of bracts in the female flowers and by

the form of the stigmas. This combination of

characters is not consistent. Finear stigmas with a

few bracts present were found in only five specimens

i.e. Van Breda 60, Muir 3060, Theron 1648, Dyer

3388 and Nelson 1 1745. In all the other specimens

the flowers are ebracteolate or only contained

weakly developed bracts. Even in plants of the

same colony ebracteolate flowers or flowers with

many or few weakly developed bracts were found.

It is therefore impossible to separate the South

African plants into different taxonomic groups.

The same conclusion was reached by Wildenauer

(MS. for Flora of S.W. Africa) when studying the

South West African material.

Apart from T. australis from Guinea several

other species of Typha have been recorded in

Africa. Their types have not been studied as they

fall outside the scope of this Flora. It is evident

YPHACEAE

55

Fig. 13.— 1, Typha latifolia subsp. capensis.fhabit ; a, inflorescence with male flowers in upper portion and

female flowers in lower portion, x ^ ( Theron 1676); b, female flower showing perianth, gynaecium and

carpodium, x 7; c, male flower showing bracteoles and stamens, X 10; d, portion ot leal showing

shape in transverse section. (1 and d from living plants at the National Botanic Garden, Pretoria;

a — c, Theron 1676).

3-F.S.A.

56

Typhaceae

that far too many species are being upheld in most

works. Examination of the dates of the existing

names indicates that the name here will not be

threatened if kept as a subspecies even if further

changes occur.

I have examined modern specimens of both

T. angustifolia and T. latifolia collected in Sweden,

and have ascertained that our plant undoubtedly

is correctly placed under T. latifolia. It differs from

typical T. latifolia in having simple, or very rarely

compound, pollen. The bracteoles of the male

flowers are brownish-white and are linear, lanceo-

late, cuneate or forked. In T. latifolia the pollen

is compound with the bracteoles of the male

flowers linear and acute.

In South Africa the plant is commonly known as

Bulrush, Catstail or Papkuil.

ORDER: HELOBIAE

by A. A. Obermeyer

The order Helobiae, also referred to as Fluviales or Najadales by some authors, consists

of a number of diverse monocotyledonous families which are usually found in or near water.

They are annuals or perennials; many are rhizomatous with the upper parts deciduous.

The leaves are sheathing at the base and mostly ligulate. An interesting feature is the presence

in all members of the minute “squamae intravaginales”, also called axillary scales, which are

thought to be secretory organs giving off mucilage. The flowers are interpreted by some as

primitive, by others as much reduced. They are usually ebracteate. The seed is exendospermous,

the embryo developing a large hypocotyl.

Engler, Wettstein, Rendle and many others, included in the Potamogetonaceae, in

addition to Potamogeton and the closely related Groenlandia, the genera here separated into

the families Zosteraceae, Ruppiaceae and Zannichelliaceae as constituted by Dandy,

Hutchinson and others. The differences in the floral structures appear to warrant this. Singh,

in his recent studies, prefers to retain Ruppia and Zannichellia in the Potamogetonaceae.

The generous assistance freely given by Mr. J. E. Dandy, Keeper of Botany, British

Museum (Natural History), is gratefully acknowledged.

Key to Families

Flowers ebracteate:

Submerged aquatics (wholly or partly) :

Carpels free, 1-6 ovulate:

Ovule solitary:

Ovule pendulous:

Flowers inside a fertile leaf-sheath on a flattened unilateral spadix,

alternately arranged in 2 rows; completely submerged grass-like

marine perennials Zosteraceae, p.57

Flowers not arranged as above:

Flowers bisexual, in spikes exserted above the water-surface during

anthesis:

Flower spikes few to many flowered, flowers arranged all around the

axis, actinomorphic, the 4 concave petals each with an opposite

sessile extrorse stamen ; drupes sessile Potamogetonaceae, p. 60

Order: Helobiae

57

Flower spike consisting of 2 naked flowers facing opposite directions

and inserted at unequal heights; the 4-8 carpels becoming

stipitate in fruit (the inflorescence then appearing umbellate);

peduncle fairly short or very long and spirally coiled. . Ruppiaceae, p. 70

Flowers unisexual, submerged, axillary, cymose or solitary, carpels 1-9,

free, sessile or stipitate in fruit; male with 2 united, apiculate

anthers (rarely 1-3) which are 6, 4, 2 or 1 locular, filament

lengthening rapidly during anthesis Zannichelliaceae, p. 73

Ovule basal, erect, anatropous; completely submerged herbs with toothed

leaves; flowers axillary, sessile, small, unisexual, often dioecious,

reduced to 1 stamen and a 1-ovuled carpel, naked or with a small

spathe Najadaceae, p. 81

Ovules 2-6 per carpel; spikes bifid (rarely 1-8) covered in bud by a calyptri-

form deciduous spathe; flowers subtended by 1-3 petaloid perianth-

segments, normally bisexual; rhizome tuberous Aponogetonaceae, p. 85

Carpels united, unilocular, with 3-6 parietal placentas, inferior, multiovulate;

flowers subtended by a bifid spathe (or 2 free bracts in the marine genus

Halophila ) Hydrocharltaceae, p. 100

Marsh plants with distichous basal leaves and ebracteate spikes;

carpels 6 (often 3 alternate sterile) connate, parting from a central axis in fruit,

uniovulate Juncaginaceae, p. 92

Flowers bracteate, with 3 sepals and 3 petals; carpels free or united, 3 — many; marsh

plants with compound umbels Alismataceae, p. 96

ZOSTERACEAE

by A. A. Obermeyer

Marine perennials, comp'etely submerged, with creeping or tuberous rhizomes, with

simple roots at the nodes. Stems, if present, floriferous, annual. Leaves distichous, convolute,

the long sheath ending in a short biauriculate ligule. Inflorescence of 1 to several unilateral

spikes each enveloped in a folded spathe which is continued above as a deciduous or semi-

persistent lamina; flowers situated in a double row on the flattened axis (“spadix”), subtended

by tongue-shaped scales (retinacula) folded inwards (these sometimes absent), monoecious or

dioecious. Male flower a solitary, sessile, unilocular, dorsifixed anther, pollen filamentous,

pollination hydrophyllous. Female flower protogynous, consisting of a unilocular ovary with

a short style and 2 long stigmatic arms; ovule solitary, pendulous, orthotropous. Fruit

a graniform nut or with 2 sagittate appendages, sometimes bursting irregularly; seed exendo-

spermous.

A widespread and common family consisting of 2 genera; one of these, the dioecious Pltyllospadix with

about 3 species, occurs on the west coast of America and in Japan, whereas the monoecious genus Zostera

with about 1 1 species is cosmopolitan.

Phillips in his Genera of South African Flowering Plants, p. 53 (1951) places this genus in the family

Potamogetonaceae .

58

ZOSTERACEAE

55

ZOSTERA

Zostera L., Sp. PI. 968 (1753); Gen. PI. ed. 5 : 415 (1754); Aschers. in Pflanzenfam. 2, 1 : 201

(1889); A. Benn. in F.C. 7 : 50 (1897); F.T.A. 8 : 225 (1901); Graebn. in Pflanzenr. 4, 2 : 27

(1907); Setchell in Proc. Nat. Acad. Sci. 19 : 810 (1933); Adamson in FI. Cape Penins. 37

(1950); Phill., Gen. ed. 2 : 53 (1951); Rendle, Class. Flow. PI. ed. 2, 1 : 207 (1953); Hutch.,

Fam. Flow. PI. ed. 2, 2 : 554 (1959); Clapham, Tutin & Warburg. FI. Brit. Isles ed. 2 : 941

(1962). Type species: Z. marina L.

Rhizome perennial, monopodial, creeping, much branched, compressed, orange-brown;

roots several from a node, simple, covered with long root-hairs. Stems, if present, floriferous,

annual. Leaves alternate, distichous, sheathing at the base; sheath tubular or convolute,

with a small biauriculate ligule; lamina linear, striate, apex rounded, bilobed or emarginate,

axillary scales 4 or 2. Inflorescences forming erect annual sympodial branches which each

terminate in a “fertile” leaf, the latter enclosing in its sheath a unilateral spadix on which

the male and female flowers are arranged alternately in 2 rows, with or without marginal

bract-like infolded lobes or “retinacula” (interpreted by some botanists as a reduced perianth);

stamen 1, sessile, unilocular, dorsifixed, pollen filiform; female consisting of a unilocular,

l-ovu!ed ovary terminating in a short style and 2 long stigmatic branches, protogynous

(the stigmas protruding from the sheath during anthesis), ovule pendulous, orthotropous.

Fruit an indehiscent nut or irregularly dehiscent, graniform, smooth or longitudinally striped

or ribbed.

Eleven species in temperate seas of the world. One species in South Africa.

Zostera capensis Setchell in Proc. Nat.

Acad. Sci. 19 : 815 (1933); Adamson in FI.

Cape Penins. 37 (1950). Type: Cape,

Knysna Lagoon, Duthie s.n. (DAV 501307,

holo.).

Z. nana sensu Krauss in Flora 344 (1845); sensu

A. Benn. in F.C. 7 : 51 (1897); sens r Graebn. in

Pflanzenr. 4, 2 : 32 (1907). Z. capensis forma

elatior Setchell, l.c. 19 : 815 (1933). Type: Knysna

Fagoon, Duthie 1212 (DAV, holo.; PRE, K).

Perennial. Roots 2-8 per node, simple,

densely covered with long root-hairs. Rhi-

zomes with internodes 5-20 mm. Stems , if

present, annual, erect, thin, bearing the axil-

lary spathes. Leaves in clusters of 2-4, con-

volute, situated on suppressed side-branches

of, and terminating the rhizomes; sheath

long, convolute, flattened below, its small

ligule forming 2 lateral rounded lobes ; lamina

linear, usually 10-30 cm (but up to 135 cm)

long, 1-2 mm broad, margin and midrib dis-

tinct, with 2-4 secondary nerves on each side

which consist of bands of smaller cells con-

taining vascular bundles alternating with

bands of larger cells containing numerous

air channels, interspersed with scattered cross-

bars, apex rounded to acute, asymmetrical,

with a narrow central slit caused by the dege-

neration of the apical cells; axillary scales

2, linear-lanceolate. Spikes on peduncles up

to 35 mm long, spathe resembling the leaf-

sheath, flattened, with the folded membranous

margins, slightly inflated in fruit, 15-30 mm

long, enclosing 3-15 flowers, the semi-persis-

tent lamina situated above the spike; retina-

cula or bract-like scales linguiform, 0-5 mm,

brown, style ultimately about 1-5 mm long

and then equalling the 2 filiform stigmata in

length. Fruit an indehiscent graniform nut,

obliquely pendulous inside the spathe, nar-

rowly ovoid, 2 mm long, yellowish brown,

closely and faintly striate, smooth. Fig 14.

Recorded from the south-west coast of South

Africa and the southern and eastern coasts as far

as Mozambique and Madagascar. Common on

muddy sandbanks in bays and estuari s, often just

exposed at low tides. Flowering occasionally

throughout the year, usually from the centre of a

colony. According to Ward’s measurements,

salinity in St. Lucia Lake varied from 18-33-5%

where Zostera occurred.

Cape. — Alexandria: Kariega River Mouth, Tolken

383. Bathurst: Bushmans River Mouth, Tolken

382; Mauve & Wells 16; Kowie Lagoon, Mauve &

Wells 6; Britten 2664; 809. Hopefield: Saldanha

Bay, Mauve 4297; Tolken s.n. Flumansdorp:

Kabeljous River Lagoon, Mauve & Wells 54.

Knysna: Lagoon, Duthie 1212; 1213; Mauve &

Wells 65 ; Plettenberg Bay, Schlechter 5935. Peddie:

Keiskama River Lagoon, Comins 1609. Kentani:

Gxaxa River, Pegler 16. Port Elizabeth: Swartkops

River Mouth, Mauve & Wells 41.

Fig. 14. — 1 , Zostera capensis, x 1; a, flowering spike (extracted from spathe), X 5; b, carpel, x 16; c, fruit

showing base of style, lateral hilum and partly exposed seed, x 16; d, leaf-tip. x 10.

60

ZOSTERACEAE

Natal. — Durban: Salisbury Island, Ward 4570.

Hlabisa: St. Lucia Lake, Ward 4172; 4175; 4294.

Lower Umfolosi: Richards Bay, Ward 4307.

Mtunzini: Umlalazi Lagoon, Ward 4019; 4252-

4256. Port Shepstone: Port Edward, lagoon, Moss

s.n. Umlazi: Isipingo Lagoon, Ward 495.

The species is closely related to the European

Z. noltii Hornern. and could perhaps be considered

a southern population as suggested by Dandy.

The two taxa are however completely isolated. The

African plants appear to be larger, the leaf apex

is asymmetrical, the retinacula are broader and the

testa of the nut is distinctly striate. Z. nana Roth

is an illegitimate name for Z. noltii Hornern.

In the Flora Capensis 7 ; 50 (1897), Bennett

enumerates a second species, Z. marina L. var.

angustifolia Hornern, based on a specimen in the

Kew Herbarium without name or collector, said

to have come from South Africa. The sheet in

question bears two identical labels “near Holyhead,

Aug. 22, 1828” and a third with CBS written on it.

Setchell believes the plants to have come from Wales.

This species has so far not been found in southern

Africa.

POTAMOGETONACEAE

by A. A. Obermeyer

Herbaceous annuals or usually perennials with deciduous stems forming colonies in

fresh or occasionally brackish water, submerged except for the spikes, which are exserted

during anthesis and, in some species, the floating leaf-blades. Roots in a circle around the

nodes of the rhizome. Rhizomes usually well developed, rarely absent, persistent, or plants

perennating by the formation of winterbuds (“turions”). Sterns erect, submerged, annual,

with the apical flower spikes just exserted. Leaves submerged and/or floating, alternate and/or

opposite, usually with a well developed convolute or tubular stipular sheath. Spikes exserted

(rarely also submerged in some North American species) few to many-flowered surrounded

by a basal sheath. Flowers ebracteate, bisexual, actinomorphic, small; perianth-segments 4;

stamens 4, sessile, extrorse, inserted on the claws of the segments; carpels 4—1, free, unilocular,

ovule solitary, campylotropous. Fruit an achene or drupe, seed exendospermous.

Cosmopolitan, comprising two genera, namely, Potamogeton with about 100 species, widespread and com-

mon, and a closely related monotypic genus Groenlandia recorded from Europe, N.W. Africa and Asia.

58 POTAMOGETON

Potamogeton L., Sp. PI. 126 (1753); Gen. PI. ed. 5 : 61 (1754); Cham. & Schlechtdl. in Linnaea

2 : 157 (1827); Aschers. in Pflanzenfam. 2, 1 : 207 (1889); A. Benn. in F.C. 7 : 45 (1897);

F.T.A. 8 : 219 (1901); Graebn. in Pflanzenr. 4, 2 ; 39 (1907); Adamson in FI. Cape Penins.

36 (1950); Phill., Gen. ed. 2 : 53 (1951); Hutch., Fam. Flow. PI. ed. 2, 2 : 556 (1959); Clapham,

Tutin & Warburg, FI. Brit. Isles ed. 2 : 942 (1962). Singh in Bot. Gaz. 126 (2) : 137-144 (1965),

morphological and anatomical studies in Helobiae. Type species: P. natans L.

Perennials, or rarely annuals, usually deciduous, surviving cold and dry periods by the

persistent woody rhizome and/or the formation of scaly winterbuds (“turions”), or in more

compatible conditions as evergreens. Roots simple or branched, several to many from a node,

often with long root-hairs. Rhizome usually well developed, much branched, sympodial,

absent or poorly developed in a few species. Stems erect, usually sparsely branched, deciduous.

Leaves often heteromorphous; in many species only submerged leaves are produced which

are thin, translucent, filiform to lanceolate, in others floating, coriaceous leaf-blades on long

petioles are formed as well; terrestrial forms from mud-banks bear coriaceous leaves only;

POTAMOGETONACEAE

61

alternate or the upper, which subtend the spikes, opposite; tissues of the lamina usually

typical for each species, with or without a lacunar band on each side of the midrib, secondary

nerves distinct, stipular sheaths convolute or tubular (the tubes usually splitting later on),

adnate to the base of the lamina (viz. in P . pectinatus and some others found outside Africa)

or free; petioles varying in length with the depth of the water in the case of the floating leaves.

Spikes pedunculate, simple, emergent (in South African species), wind- or rarely water-

pollinated ( P . pectinatus ), few- to many-flowered, ovoid to cylindric, dense, lax or interrupted,

lengthening during anthesis and fruiting. Flowers ebracteate, bisexual, actinomorphic,

protogynous, small; perianth-segments 4, free, clawed, concave; stamens sessile, extrorse,

straddling the claw of the segment, pollen globose; carpels 4, occasionally fewer, ovoid,

with a beak bearing the stigma or the stigmatic area extended on the ventral suture; ovule

solitary, laterally attached. Fruit an ovoid drupe with a short or a long beak, a soft epicarp

and a hard endocarp; cotyledon bent like a hook, with a large hypocotyl.

A cosmopolitan genus containing about 100 species with 7 in Southern Africa.

The floral parts have been variously interpreted. Graebner, Rendle and others believed the “petals” to

be appendages to the connective of the anther. Uhl's studies in 1947 and various subsequent botanists

.support Kunth’s theory (1841) that the flower actually represents an inflorescence containing male flowers,

each surrounded by a petaloid bract, and naked female flowers.

Gender of the generic names Potamogeton and Aponogeton: — “Both were regarded as neuter by their

original authors but are now generally treated as masculine as in the Pflanzenreich and other works. The

Greek word geiton (a neighbour) can be either masculine or feminine but not neuter. In view of the fact that

the masculine gender has been adopted for them both, I see no reason to interfere with this view” — J. E.

Dandy in letter dated 2.8.1963.

Leaves all submerged, narrow; stipules adnate to the lamina below where it forms a folded sheath with

a whitish margin, free as a ligule above; spikes few-flowered, interrupted; stigma with large papil-

lae, water pollinated (Subgenus Coleogeton) 1 . P. pectinatus

Leaves either all submerged or also with floating blades variously shaped; stipules free from the leaf,

folded or tubular; spikes usually not interrupted; stigma with small papillae, wind pollinated

(Subgenus Potamogeton ):

Submerged leaves very narrow 0-5^4 mm wide:

Leaves all submerged; lacunar system bordering midrib not developed or very narrow:

Stipule tubular at least when young, pale brown; leaves usually about 2 mm wide; fruit smooth

2. P. pusillus

Stipule folded; leaves usually 1 mm wide; fruit usually tuberculate and with a basal tooth

3. P. trichoides

Leaves submerged but with floating involucral leaves subtending the flower spikes; submerged

leaves with a well developed lacunar system bordering the midrib; stipule folded ... .4. P. octandrus

Submerged leaves over 4 mm wide or plants terrestrial :

Leaves all submerged, uniform:

Leaf margin toothed and undulate; spikes few-flowered; beak nearly as long as the fruit

5. P. crisp us

Leaf margin entire, smooth or undulate, upper leaves sometimes floating, net-veined; spikes

many-flowered ; beak short 6 . P. schweinfurthii

Primary submerged leaves if in deep water soon disintegrating; mature leaves floating with an

oblong, coriaceous lamina, margin smooth, veining indistinct; plants often creeping on mud-

banks, exposed 7. P. thunbergii

1. Potamogeton pectinatus L., Sp.

PI. 127 (1753); A. Rich., Tent. FI. Abyss.

2 : 354 (1851); A. Benn. in J. Bot. Lond.

33 ; 138 (1895); F.C. 7 : 49 (1897); F.T.A.

8 : 223 (1901), pro parte; Dur. & Schinz,

Consp. FI. Afr. 5 : 495 (1894); Graebn. in

Pflanzenr. 4, 2 : 121, fig. 28 A-B (1907), pro

parte; Dinter in Fedde, Rep. 22 : 383 (1926);

Dandy in Journ. Linn. Soc. Bot. 50 : 513

(1937); Adamson in FI. Cape Penins. 36

(1950). Type: from Europe.

P. filiforme sensu A. Benn. in F.T.A. 8 : 223

(1901); P. livingstonei A. Benn. l.c. (1901). Syn-

types: Malawi, Lake Nyasa, Kirk s.n.; Laws 3;

Bellingham s.n. (BM, K). P. pusillus sensu Marloth,

FI. S. Afr. 4 : 13, fig. IV, 1, 5, 6 (1915). P. pectinatus

var. ungulatus Hagstr. in Svensk. Vet. Akad.

62

POTAMOGETONACEAE

Fig ] 5 — i Potamogeton pectinatus, x 1; la, fruits, X 4; lb, lc, leaf-base with attached sheath, x 4,

Id, winterbud, x 3. 2, P. trichoides, x 1; 2a, leaf apex, X 15; 2b, free, open leaf-sheath, x 8; 2c,

fruit, x 1.

POTAMOGETONACEAE

63

Handl. n.s. 55 : 39 (1916). Type: Cape, Caledon

District, Koude River, 1896, Schlechter 9592

(S, holo., PRE). Hagstrom does not mention a

number but it is presumably this gathering.

Submerged perennials forming large

colonies, found in fresh or brackish water,

the upper parts dying down in cold or dry

spells. Rhizome persistent, forming scaly

winter buds (“turions”), much branched, with

numerous long fine roots arising in whorls

from the nodes, the roots bearing long root-

hairs. Stems varying in length in proportion

to depth of the water, branching freely, terete,

slender. Leaves all submerged, often bunched

above and spreading in a fan-shaped manner;

stipules attached to the lamina, forming a

folded sheath, often with whitish margins and

terminating above in a large, rounded deci-

duous ligule; lamina usually curved ou . wards

above the sheath, linear to filiform, 4-12 cm

long and 1-2 mm broad, translucent, with

cross-bars, if broad with lateral longitudinal

veins, apex acute or rounded and apiculate.

Spikes lengthening during anthesis and the

fruiting stage, the lower flowers especially

wide'y spaced, scape wiry, thin, whorls 4-8,

consisting of 2-4 flowers each, stigmas large

(the pollen or detached flowers floating on

the water and thus contacting thelarge papillae

on the stigmas). Fruit c. 4 mm long, yellowish-

brown, convex, dorsally with 2 distinct broad

lateral keels sloping inwards to the ventral

straight keel, beak short. Fig. 15 : 1.

More or less cosmopolitan. Common throughout

the Republic and South West Africa in brackish

and fresh water.

Cape. — Bredasdorp: Hagel, Kraal River, Leighton

2521. Caledon: Koude River, near Elim, Schlechter

9592. George: Swartvlei, Chunnett s.n. Kentani:

Gxaxa River, Pegler 438. Kimberley: Riverton,

Tapscott s.n. Namaqualand: Steinkopf, Meyer sub

Marloth 6465. Peninsula: Prince’s Vlei, Marloth

7288; Zeekoevlei, Adamson 3580. Port Elizabeth:

Swartkops River, Ecklon & Zeyher 644. Robertson:

Breede River, Koch s.n.

Natal. — Durban: Umgeni River Mouth,

Schlechter 2969. Hlabisa: St. Lucia Lake, False

Bay, Ward 4280. Ingwavuma: Kosi Estuary,

Tinley 286.

O.F.S. — Boshof: Smitskraal, Burtt Davy 1 1332.

Kroonstad: Vais River, Pont 599.

Transvaal. — Ermelo: Lake Chrissie, Moss 16293.

Germiston: Lake, Burtt Davy 2534. Potchefstroom :

Vaal River, Louw 1634. Wolmaransstad : Liebenberg

2456.

S.W.A. — Grootfontein: Grosshuis, Schoenfelder

1008. Rehoboth: Buellsport, Rodin 2973.

This is the only South African species of Potamo-

geton tolerating brackish water. In Zululand at

Lake St. Lucia, Ward measured patches of 45 x 32

feet with the adult plants mainly peripheral. Where

they occurred salinity varied from 19 to 26-5%

according to his tests. Occasionally the plants

are encrusted with lime.

2. Potamogeton pusillus L., Sp. PI. 127

(1753); Dandy & Taylor in J. Bot. Lond.

76 : 90 (1838); Clapham, Tutin & Warburg,

FI. Brit. Isles, ed. 2 : 953 (1962). Type:

Europe (LINN 175-15, left hand specimen,

lecto.).

P. panonnitanus Biv., Nouv. Pi. 6 (1838); Hagstr.

in Svensk. Vet. Akad. Handl. n.s. 55 : 98 (1916);

Dandy in Journ. Linn. Soc. Bot. 50 : 523 (1937).

Type: Europe, Sicily. P. friesii sensu A. Benn. in

F.C. 7 : 48 (1897). P. preussii sensu A. Benn.

in F.T.A. 8 : 222 (1901). P. pusillus var. africanus

A. Benn. in Ann. Cons. Jard. Geneve 9 : 102 (1905);

Graebn. in Pflanzenr. 4, 2 : 115 (1907). Syntypes:

Transvaal, Wilms 1656; 1657 (G); Natal, Umlazi

River, Drege 1206 (G); Umzinto, Wood 3055

(G, K, NH, BOL). P. mucronatus sensu Graebn.

in Pflanzenr. 4, 2 : 113 (1907). P. badius Hagstr.

in Svensk. Vet. Akad. Handl. n.s. 55 : 104 (1916).

Type: Natal, Umlazi, Drege 4458 (S, holo.).

P. exiguus Hagstr. in Svensk. Vet. Akad. Handl.

n.s. 55 : 106, fig. 42 (1916). Type: Cape Town, 53,

Andersson (S, holo.). P. subjavanieus Hagstr. in

Svensk. Vet. Akad. Handl. n.s. 55 : 129 (1916).

Type: Cape, Swartkops River, 1830, prob. Zeyher

(S, holo., PRE, photo.).

Submerged herbs arising from winter-

buds or seed and forming no rhizome or a

poorly developed one. Roots long and fine.

Stems up to 100 cm, branching from near the

base, internodes fairly equal in length, thin,

slightly striate and compressed. Leaves all

submerged, firm, translucent, linear, usually

c. 20-40 mm long and 1-2 mm broad, apex

obtuse, with 2 lateral longitudinal veins

joining the midrib abruptly near the tip at

unequal heights; areas between the lateral

and central veins without a distinct lacunar

system, sometimes with a narrow dense band

of small cells flanking the midrib; stipular

sheaths 6-17 mm, tubular, splitting later,

usually semi-persistent, pale brown. Spikes

axillary, usually shorter than the leaves;

scape thin, becoming firm in fruit, 1-4 cm

long; flowers 2-4 per whorl with up to 4

whorls per spike. Drupe 2-2-5 mm long and

1-1-5 mm broad, pale olive green, obovoid,

smooth, ventral face convex, dorsal more

strongly convex, with a faint brown keel

when dry, beak short and straight. Fig. 16:2.

64

POTAMOGETONACEAE

Fig. 16. — 1, Potamogeton octandrus, habit, x 1; la, free, open leaf-sheath, x 4; lb, drupe in longitudinal

section showing hooked embryo; 1c, lamina showing lacunar tissue beside midrib. 2, P. pusillus, habit,

x 1; 2a, drupe, x 8; 2b, free, tubular leaf-sheath, x 3 ; 2c, leaf apex showing insertion of lateral nerves,

X 7; 2d, lamina showing raised midrib and dense tissue beside midrib, X 9.

POT A MOG ETONACEAE

65

Widespread in the northern hemisphere and

Africa. Fairly common in Southern Africa; occurs

in fresh water.

Cape. — Alexandria: de Bega River, Archibald

4206. Bathurst: Grahamstown, MacOwan 1992.

Caledon: Stormsvallei, Riviersondereinde, Zeyher

1733 (BOL). Calvinia: Lokenburg, near Nieuwoudt-

ville, Story 4419. King William’sTown: Yellowwoods

River, Flanagan 2189. Komga: near Komga,

Flanagan 2374. Mqanduli: Qingqolo, Pegler 2042.

Mount Currie: Kokstad, Tyson 1807. Port Elizabeth :

Happy Valley, Long 1 145. Port St. Johns: Umgazi

River, Bolus 10374. Uitenhage: Swartkops River,

Mauve & {Veils 55.

Natal. — Bergville: Cathedral Peak Forest

Reserve, Inhlwasine River, Killick 1608. Camper-

down: near Camperdown, Rehmann 7686. Durban:

Umbilo, Moss 5449. Hlabisa: Hluhluwe Game

Reserve, Ward 2505; 2576. Inanda: Umhlangwe

River near Phoenix, Schlechter 3122. Umzinto:

Dumisa, Rudatis 2139.

Basutoland. — Between Bokong and Modiba

Matso Rivers, Coetzee 519. Leribe: Hlotse River,

Makokoane, Dieterlen 1033.

O.F.S. — Bloemfontein: Glen, Modder River,

Wager s.n.

Transvaal. — Johannesburg: Moss 9681. Lyden-

burg: Wilms, s.n. Marico: Wonderfontein, Burtt

Davy 7610. Petersburg: Houtbos, Schlechter

4405; Turfloop, Schlechter 4695. Potchefstroom:

Louw 1614. Pretoria: Rietvlei Dam, Mauve 4259.

Swaziland. — Mankaiana: Hlotane River,

Compton 30876.

S.W.A. — Warmbad: Fish River Canyon, Comins

1795.

In a few representatives some of the younger

upper leaves form a narrow spathulate floating

lamina and they would therefore resort under the

species P. subjavanicus Hagstr. Dandy suggests

that this last-named species is a form of P. pusillus

L. and he places the following gatherings under it;

Natal. — Eshowe: Gingindhlovu, Gerstner in NH

22187. Inanda: near Verulam, Wood 962. Cape. —

Vanrhynsdorp : Klaver, Smith 2602.

In appearance the plants resemble the tropical

species P. octandrus Poir. but the tubular leaf-

sheath of P. subjavanicus Hagstr. separates it from

that species. So far no fruiting material has been

collected.

3. Potamogeton trichoides Cham. & Sch-

iechtd. in Linnaea 2 : 175, t.4, fig. 6 (1827);

Graebn. in Pflanzenr. 4, 2 : 119(1 907) ; Hagstr.

in Svensk. Vet. Akad. Handl. n.s. 55 : 74, fig.

29 (1916); Dandy in Journ. Linn. Soc. Bot.

50 : 521 (1937); Clapham, Tu'in & Warburg,

FI. Brit. Isles ed. 2 : 955 (1962). Type: Europe.

Small wiry herbs forming dense masses,

50-75 cm high. Rhizome poorly developed or

absent. Stems thin, 0-25 mm in diarn., groov-

ed, much branched. Leaves all submerged,

linear to setaceous, c.4 cm long and 1 mm

broad, translucent, spreading, flat, apex

acuminate, midrib prominent, yellowish;

stipular sheaths open, deciduous, c.8 mm

long, membranous. Spikes very small, shorter

than the leaves, c.4-flowered, stalk wiry, often

curved above; petals with long claws, carpel

1, rarely more, with a large peltate stigma.

Drupe ovoid, c.l mm long, ventral side nearly

straight, beak short, sometimes tuberculate.

Fig. 15:2.

Found in Europe, western Asia and Africa.

Widespread but apparently rare in Southern Africa;

in fresh water ponds.

Cape. — Albany: near Grahamstown, Slaaikraal,

Mauve & Wells 18. Uitenhage: Redhouse, Holland

4169 (GRA).

Natal. — Dundee: Verdrukdam, Hattingspruit,

Edwards 3132.

Transvaal. — Johannesburg: Tonquani, 12 miles

N.W. of Johannesburg, Mogg 23777.

4. Potamogeton octandrus Poir. in Lam.,

Encycl. Meth. Bot., Suppl. 4 : 534 (1816).

Hutch, in F.W.T.A. 2 : 307 (1836); Dandy

in Journ. Linn. Soc. 50 : 517 (1937). Type:

Cochinchina, Loureiro (BM, holo.).

Hvdrogeton heterophylhis Lour., FI. Cochinch.

1 : 244 (1790), non Potamogeton heterophyUus

Schreb. (1771). Type: Cochinchina.

Potamogeton javanicus Hassk. in Act. Soc. Sci.

Ind.-Neer). 1, 8 : 26 (1856); A. Benn. in Journ.

Bot. Lond. 29 : 121, 186 (1891); l.c. 33 : 138 (1895);

F.T.A. 8 : 220 (1901); Dur. & Schinz, Consp. FI.

Afr. 5 : 495 (1894); Graebn. in Pflanzenr. 4, 2 : 46,

fig. 14 A-C (1907). Type: Indonesia.

Herbs 20-100 cm high from a thin

creeping rhizome, forming slender winter-

buds. Stems with long internodes below,

short above, grooved, producing numerous

fairly short side branches above. Leaves

dimorphous; submerged leaves alternate,

translucent, soft, linear to narrowly linear-

lanceolate, 3-7 cm long and 1-3 mm broad,

apex acute or acuminate, the area between

the midrib and lateral nerves consisting of

lacunar, reticulated tissue with larger cells,

the lateral nerves becoming faint towards the

tip, approaching the midrib gradually; upper

floating “involucral” leaves opposite, just

below the flowering spikes, lanceolate to

oblong-lanceolate, up to 2 cm long and 5 mm

broad, narrowed into a petiole of 1 cm long,

thinly coriaceous, not transparent, apple

green, with about 7 nerves showing up dis-

tinctly below (some leaves showing transi-

tional stages from the submerged to the

66

POTAMOGETONACEAE

floating forms); stipular sheaths convolute,

small, soon decaying, upper somewhat larger

and broader. Spikes small, 25-40 mm long,

lengthening during fruiting, whorls 3-7, each

whorl usually 2-flowered, scape flattened,

slightly thicker above, about 15 mm long.

Fruit about 2 mm long and 1 • 5 mm broad,

rounded to oblong in outline, broad-shoul-

dered above, tapered to the base, ventral keel

nearly straight, with a central tooth, dorsal

keel crenulate ending in a distinct tooth,

lateral keels convex ending in teeth near the

base, beak fairly long and broad, upturned.

Fig. 16 : 1.

Widely distributed in tropical Africa, Asia and

Australia. So far recorded only from northern and

eastern parts of Southern Africa.

Natal. — Inanda: Umhlanga River near Phoenix,

Schlechter 3125. Mtunzini: near Amatikulu, Ward

2560. New Hanover: York, Umshati River, Wood

4300.

Transvaal. — Johannesburg: Florida Lake,

W'ager J. 35945. Pretoria: Pienaars River, Mauve &

Schlieben 9598. Waterberg: Badsloop, Schlechter

4780. Witbank: near Witbank, Repton 952.

S.W.A. — Okavango Native Territory: Okavango

River at Runtu, De Winter & Marais 4477; Merx-

mii/Ier & Giess 1891. Ovamboland: Kilevi on the

Kunene River, Schinz 1001.

When in full flower the spikes appear yellow

because of the copious pollen.

5. Potamogeton crispus L., Sp. PI. 126

(1753); Solms in Beitr. FI. Aethiop. 1 : 194

(1867); Dur. & Schinz, Consp. FI. Afr. 5 : 493

(1894). pro parte; A. Benn. in Joum. Bot.

Lond. 33 : 137 (1895); F.T.A. 8 : 221 (1901);

Graebn. in Pflanzenr. 4, 2 : 97 fig. 23 (1907);

Dandy in Journ. Linn. Soc. Bot. 50 : 537

(1937); Clapham, Tutin & Warburg, FI.

Brit. Isles ed. 2, 956 (1962). Type: Europe

(LINN 175-6).

P. crispus L. var. najadoides Graebn. in Pflanzenr.

4, 2 : 100 (1907). Type: Transvaal, Standerton,

Wilms 1658 (Bf).

Fairly large plants 50-150 cm high, with

much ramified perennating rhizomes which

also form horny winterbuds. Stems slender,

4-angled, branched above. Leaves all sub-

merged, sessile, linear-lanceolate, 3-9 cm

long and 8-15 mm broad, apex obtuse, apicu-

late, margin finely serrate, markedly undulate,

often reddish, shiny ranslucent, longitudinal

veins 3-5, the laterals close to the margin;

stipular shea'.h convo’ute, 10-20 mm, sub-

triangular, obtuse, early deciduous. Spikes

few-flowered, somewhat lax, peduncle 2-5-7

cm long, slender, narrowed upwards, usually

curved; flowers 5-10. Drupe 2-4 mm long,

ovoid-acuminate, laterally compressed, with

a prominent pectinate dorsal keel, the ventral

keel fairly straight, beak nearly as long as the

fruit, falcate, tapered. Fig. 17.

Recorded from Europe, Asia, Africa and

apparently introduced in America. Fairly common

in fresh water in summer rainfall areas of Southern

Africa.

Cape. — Barkly West: Vaal River at Sydney,

Acocks & Haf strom 862. Komga: Flanagan s.n.

Mount Currie: Clydesdale, Umzimkulu River,

Tyson 2125. Prieska: Orange River, Bryant 733.

Taungs: Harts River, Parker.

O. F.S. — Bloemfontein: Glen, Modder River,

Mostert 892. Boshof: Smitskraal, Vaal River,

Burtt Davy 21. Kroonstad: Vais River, Pont 598.

Natal. — Camperdown: Wood 11107. Hlabisa:

Hluhluwe Game Reserve, Ward 2667. Ingwavuma:

Mahlangwe Pan, Tinley 222. Umlazi: Umlaas

Wood 154.

Transvaal. — Brits: Assen, junction of Pienaars

and Crocodile Rivers, Obermeyer in TRV 34714.

Pilgrim’s Rest : Kruger National Park, Olifants River,

Codd 6193. Pretoria: Rust de Winter Dam, Wouda

1. Vereeniging: Wager in TRV 10455.

Clos in France (Bull. Soc. Bot. France 3, 1856),

in his researches on the peculiar propagation of

P. crispus, drew attention to the unique horny

turions. The detached compressed shoot forming

the turion grows no further, but its axillary buds

form new plants.

6. Potamogeton schweinfurthii A. Benn.

in F.T.A. 8 : 220 (1901); Graebn. in Pflanzenr.

4, 2 : 79, fig. 19 (1907); Hutch, in F.W.T.A.

2 : 307 (1936); Dandy in Journ. Linn. Soc.

Bot. 50 : 526 (1937). Type: Sudan, Upper

Nile Province, Schweinfurth 1223. (K, sheet

1, lecto.; NH).

P. lucens var. fluitans sensu A. Benn. in F.C.

7 : 48 (1897); F.T.A. 8 : 221 (1901). P. capensis

Scheele ex Hagstr. in Svensk. Vet. Akad. Handl.

n.s. 55 : 203, fig. 101 (1916). Syntypes: Cape,

Uitenhage, Swartkops River, Zeyher (S, 919, 8799);

Natal, Umshlangwe River, Schlechter 3120 (S).

P. promontoricus Hagstr. in Svensk. Vet. Adad.

Handl. n.s. 55 : 182, fig. 94 (1916). Type: Cap.

Bonae Spei, Drege (UPS). P. venosus A. Benn. in

Trans. & Proc. Bot. Soc. Edin. 29 : 52 (1924).

Type: Natal, Umhlongwe, Wood 3015 (K, holo;

NH, BOL).

Large submerged plants up to 200 cm

long from a perennial creeping woody rhi-

zome with internodes 4—8 cm long. Stems

erect 30-200 cm high, 1-3 mm in diam.,

terete. Leaves submerged or occasionally

POTAMOGETONACEAE

67

the upper floating, sessile, linear-lanceolate,

attenuated to the base and apex, 10-20 cm

long and 1-2 cm broad, apex apiculate, mar-

gin smooth, finely undulate, distinctly net-

veined with the cross-veins oblique, ascend-

ing, midrib prominent, lamina of the sub-

merged leaves thin and soft, the exposed ones

coriaceous, shiny; stipules persistent, folded,

the lower amplexicaul, the upper spreading,

large, lanceolate, 2-4 cm long. Spikes on

swollen scapes, 4-10 cm long, many flowered.

Drupe ovoid, 3-4 mm, smooth, ventral side

fairly straight, dorsal rounded, beak short,

occasionally with lateral keels. Fig. 18.

Widespread in northern and eastern tropical

Africa extending southwards to the eastern Cape,

Natal, Transvaal and South West Africa. Common in

rivers in the Transvaal.

Cape. — Kimberley: Danielskuil, Wilmati 1252.

Uitenhage: Swartkops River, Ecklon & Zeyher 640.

Transvaal. — Klerksdorp: Schoonspruit near

Klerksdorp, Nelson 224. Letaba: Kruger National

Park, Rooi Rotse, Van der Schijff 5679. Potchef-

stroom: Vaal River at Schoemansdrift, Louw 1512.

Pretoria: Rietvlei Dam, Mauve 4257; Rust de

Winter Dam, Wouda 3. Vereeniging: Vaal River,

Moss 13372.

Natal. — Hlabisa: Hluhluwe Game Reserve,

Ward 2567 ; 2577; 4203. Ingwavuma: Lake.

Sibayi, Tinley 67.

S.W.A. — Grootfontein : Otavi, Dinter 5254*

Rehoboth: Buellsport, Strey 2004; Rodin 297 5.

7. Potamogeton thunbergii Cham. & Sch-

lechtci. in Linnaea 2 : 221, t.6, fig. 21 (1827).

Type: Cape, Swellendam, Hartebeeskraal

near Brak River, Mund & Maire (Bf, holo.).

Fig. 17 — 1, Potamogeton crispus, x 1 ; a, overwintering side shoot, x 1 ; b, drupe, x 5; c, leaf apex,

showing teeth and nerves, x 5.

68

POTAMOGETONACEAE

Fig. 18. — 1, Potamogeton schweinfurthii, x 1; a, submerged sessile leaf, X 1; b, drupe, x 9.

POTAMOGETONACEAE

69

Fig. 19. — 1, Potamogeton thunbergii, X 1; a, submerged petioled leaf, X 1; b, young sprouting shoots from

winterbuds, x 1 ; c, diagrammatic figure of flower showing perianth segments, part of stamens and 2

carpels, x 6; d, young drupe, x 6; e, mature drupe, x 4.

70

POTAMOGETONACEAE

P. capensis Mund & Maire ex Cham. & Schlechtd.

i.c. (The name P. capensis Scheele nom. nud. in

herb. Buchenau, Bremen, was quoted in error under

this species by Bennett in F.C. 7 : 46 (1897); it

belongs to P. schweinfurthii). P. natans sensu

Thunb., Prodr. 32 (1794); sensu A. Benn. in F.C.

7 : 46 (1897). P. natans var. capensis sensu Dur. &

Schinz, Consp. FI. Afr. 5 : 494 (1895). P. richardii

Solms in Schweinf., Beitr. FI. Aethiop. 194, 292

(1867); Dur. & Schinz, Consp. FI. Afr. 5 : 496

(1895); A. Benn. in F.C. 7 : 47 (1897); F.T.A.

8 : 220 (1901); Graebn. in Pflanzenr. 4, 2 : 56

(1907); Hagstr. in Svensk. Vet. Akad. Flandl.

n.s. 55 : 104 (1916); Wild in Kirkia 2 : 20, t.8

(1961). Type: Ethiopia, Schimper 135 (BM, K).

P. fluitans sensu A. Benn. in F.C. 7 : 46 (1897).

P. fluitans subsp. americanus proles thunbergii

sensu Graebn. in Pflanzenr. 4, 2 : 61 (1907). P.

americanus var. thunbergii sensu A. Benn. in F.C.

7 : 47 (1897). ? P. fibrosus Hagstr. in Svensk. Vet.

Akad. Handl. n.s. 55 : 160 (1916). Type: Without

locality or collector, 91 (S, holo.). Dandy provi-

sionally includes it under this species.

Perennial with rhizomes much ramified,

matted, woody, usually orange-brown, terete,

with the internodes up to 8 cm long; scaly

winterbuds formed in autumn. Stems erect,

usually unbranched, up to 100 cm long.

Leaves with submerged and floating forms

or, if on land, with aerial blades; in deep

water the submerged (juvenile) leaves decay

before or during flowering; all the leaves

petioled, the length varying with the depth

of the water; lamina of the submerged leaves

linear to linear-lanceolate, tapering to the

base and apex, 8-20 cm long and 8-16 mm

broad, thin, net-veined, with the transverse

veins oblique, ascending; floating lamina

elliptic to ovate, obtuse to rounded at the

base, apex rounded to acute, about 5 cm long

and 2-5 cm broad (up to 8 x 3-5 cm) coria-

ceous, veins indistinct; intermediate forms

between the submerged and floating forms

occur; leaves of land forms usually smaller,

midrib prominent below; stipules folded,

membranous, lower amplexicaul, upper

spreading, ovate, acuminate, variable in size,

up to 6 cm long. Spikes on somewhat swollen

arcuate scapes, 5-10 cm long, flowering part

cylindrical up to 5 cm long in fruit, many

flowered. Drupe rounded, compressed c.4 mm

long, green to brown, the convex dorsal side

3-carinate, the keels terminating in 3 basal

knobs, sides loping inwards to a ventral

keel which is indented or straight. Fig. 19.

Distributed from east tropical Africa to South

West Africa and the eastern Cape; common in

the Transvaal. Grows on mudbanks and in deep

water.

Cape. — Albany: Vlei 4 miles E. of Grahamstown,

Britten 5033. Alexandria: near Bushmans River,

Comins 1846. Knysna: Portland, Duthie 1172.

Prince Albert: Swartberg, Pocock S.232.

Basutoland. — Leribe: near Hlotse, Dieterlen 831.

Natal. — Bergville: Edwards 2408. Durban:

Bluff, Huntley 135. Hlabisa: Makhakhatana,

Vincent, Ward & Wells 5. Umvoti: Edwards 2752.

Transvaal.— Boksburg: Codd 6114. Letaba:

Westfalia Estate, Scheepers 849. Middelburg:

Strey & Schlieben 3836. Potchefstroom: Leeu-

fontein, Louw 1720. Pretoria: Pienaars River,

Mauve & Schlieben 9595.

S.W.A. — Caprivi: Singalamwe, Killick & Leistner

3229. Grootfontein North: Runtu, Okavango

River, Merxmiiller & Giess 1889.

Closely related to P. nodosus Poir. from Europe

and N.W. Africa, but distinguished from it by its

smaller leaves (which attain only half the size of

the European species) the drupe which has well

developed knobs on the dorsal side and a straight

ventral keel, etc.

RUPPIACEAE

by A. A. Obermeyer

Submerged slender grasslike herbs found in saline pools near the sea. Rhizome matted,

perennial. Stems erect, thin. Leaves setaceous, sheathing at the base. Inflorescence on a short

or long spirally coiled peduncle, flowers 2 on each spike, bisexual, at unequal heights and

opposite sides of the rhachis, just exserted above the water at anthesis, stamens 2, with divergent

locules, pollen luniform; carpels 4 or more, becoming long stipitate in fruit.

Monogeneric.

Ruppiaceae

71

59 RUPPIA

Ruppia L., Sp. PI. 127 (1753); Gen. PI. ed. 5 : 61 (1754); Aschers. in Pflanzenfam. 2, 1 : 210

(1889); A. Benn. in F.C. 7 : 49 (1897); F.T.A. 8 : 224 (1901); Graebn. in Pflanzenr. 4, 2 : 142

(1907); Adamson in FI. Cape Penins. 36 (1950); Phill., Gen. ed. 2 : 54 (1951); Hutch.,

Fam. Flow. PI. ed. 2, 2 : 558 (1959); Clapham, Tutin & Warburg, FI. Brit. Isles ed. 2 : 959

(1962). Singh in Bot. Gaz. 126 (2) : 138 (1965). Type species: R. marina L.

Perennial herbs completely submerged, forming large colonies with branches up to about

75 cm long. Roots simple, 1-2 per node, densely covered with very long root-hairs. Rhizomes

creeping, branched, thin, internodes angled. Stems thin, branching freely. Leaves alternate

or opposite above, spreading, lamina linear to setaceous, up to 10 cm long and 1 mm broad,

stipular sheaths attached to the base of the lamina, wide, folded, membranous, obtuse at the

apex, amplexicaul at the base. Inflorescences at first enclosed in the swollen sheaths of the

apical leaves, consisting of 2 opposite flowers inserted at different heights, viz. spicate but

appearing sub-umbellate in fruit because of the stipitate drupes; peduncle remaining short

in one species, accrescent in the other, becoming long and spirally coiled. Flowers bisexual,

naked, protandrous; stamens 2, opposite, the separated locules divergent with their short

filiform connectives attached to the short swollen columnar filament, pollen luniform, buoyant ;

carpels usually 4, occasionally more, sessile at first, becoming long-stipitate in fruit, ovule

solitary, pendulous, campylotropous, stigma sessile, peltate. Fruits indehiscent on stipes 1-3

cm long, pyriform, often oblique, beaked, sometimes verruculose; seed hard, testa brown.

Two species recognised which are found in temperate and subtropical regions in brackish water near

the sea or, rarely, in mountain lakes in the tropics. (According to some authors the genus can be divided

into 10-15 species).

Phillips in his Genera of South African Flowering Plants p. 54 (1951) places this genus in the family

Potamogetonaceae.

Fruiting scape usually short and recurved, occasionally up to twice as long as fruiting stipes; leaves

acute 1. R. maritima

Fruiting scape long and spirally twisted; leaves obtuse or rounded 2. R. spiralis

1. Ruppia maritima L., Sp. PI. 127 (1753);

Aschers. in Pflanzenfam. 2, 1 : 210 (1889);

Rendle in Cat. Afr. PI. Welw. 94 (1899);

Graebn. in Pflanzenr. 4, 2 : 142 (1907).

Adamson in FI. Cape Penins. 36 (1950). Type:

Europe (LINN 1761).

R. rostellata Koch in Reichb., Icon. PI. Crit.

2 : 66, 1. 1 74, fig. 306 (1824); A. Benn. in F.T.A.

8 : 224 (1902). Type: Europe.

Leaves acute; scape usually short and

straight not more than about twice as long

as the fruiting pedicels; otherwise characters

of the genus. Chromosome number 2n = 20.

Fig. 20 : 2.

Cosmopolitan; common in brackish water with

weak salinity along the coast from South West

Africa to Zululand.

Cape. — East London: Hagahaga Mouth, Flanagan

2366. Knysna: Groenvlei, Martin 4452. Peninsula:

Maitland Bridge, Wolley Dod 3644; Noordhoek

Flats, Mauve 4254.

Natal. — Ingwavuma: Ndumu Game Reserve

Tinley 507.

S.W.A. — Swakopmund: Swakop River Mouth,

Merxmiiller & Giess 1739.

2. Ruppia spiralis L. ex Dum., FI. Belg.

164 (1827); A. Benn. in F.C. 7 : 50 (1897);

F.T.A. 7 : 224 (1901); Clapham, Tutin &

Warburg, FI. Brit. Isles ed. 2 : 959 (1962).

Type: Europe (LINN 176-2).

R. maritima L. subsp. spiralis (L. ex Dum.)

Graebn. in Pflanzenr. 4, 2 : 142 (1907).

Leaves obtuse or rounded; scape long,

spirally coiled, the flowers just exserted dur-

ing anthesis; otherwise characters of the

genus. Chromosome number: 2n=40. Fig

20 : 1.

Cosmopolitan but rare in the tropics. Needs a

more saline medium than R. maritima. Found along

the coast from Namaqualand to northern Natal.

72

Ruppiaceae

Cape.— Bathurst: Port Alfred, Saltvlei, Britten

2694. Clan william: Lamberts Bay, Langevlei,

Tolken 335. George: Swartvlei, Mauve & Wells 68.

Namaqualand: Hondeklip Bay, Pillans in BOL

18256. Peninsula: Kommetjie Effluence, Wolley

Dod 3472; Noordhoek salt pan, Tolken 444;

Rugby, Tolken 449. Port Elizabeth: Redhouse,

Holland 4449.

Natal. — Hlabisa: Lake St. Lucia, Ward 4173;

4176; 4178; 4180; 4295.

Fig. 20.— 1, Ruppia spiralis, X 1 ; la, flowering spike with ripe anthers, lb, flowering spike with shrivelled

anthers and ripe carpels, x 10; lc, leaf-tip, x 12; Id, drupe, x 10. 2, R. maritima, x 1; 2a, leaf-tip,

X 12; 2b, drupe, x 10.

Zannichelliaceae

ZANNICHELLIACEAE

by A. A. Obermeyer

73

Submerged annual or perennial herbs with the leafy stems permanent or deciduous.

Roots adventitious appearing from an annular skinfold at the rhizome-nodes. Rhizomes

creeping, matted, bracteate. Leaves alternate, opposite or ternate, often congested above,

setaceous, linear, strap-shaped or tubular, the stipules adnate to the base of the lamina or free-

forming a folded or tubular sheath; those of the upper fertile leaves modified into bracts

with the lamina reduced or wanting. Flowers unisexual, monoecious or dioecious, solitary or

cymose; “perianth” absent or present, flask-shaped or rarely of 3 small scales; male with 2

(rarely 1 or 3) united anthers, 6, 4, 2 or 1 locular, with an apical appendage, filament (or pedicel)

lengthening rapidly at anthesis, pollen globose or filamentous, pollination hydrophyllous;

carpels free, style long or short, simple, stigma capitate, peltate, funnel-shaped, or 2-4 lobed,

ovule solitary, pendulous. Fruit a drupelet with the cotyledon spirally coiled, exendospermous;

rarely viviparous ( Amphibolis antarctica ).

Cosmopolitan; 4 marine genera and 3 others which inhabit brackish or fresh water. Four genera recorded

in Southern Africa, 2 of these marine.

Phillips in his Genera of South African Flowering Plants p. 54 (1951) places the genera Zannichellia and

Althenia in the family Potamogetonaceae. The other two genera had not yet been recorded for South

Africa in 1951 but were also placed in the family Potamogetonaceae by Graebner in Das Pflanzenreich 4, 2

(1907).

Marine plants; dioecious; pollen filamentous:

Leaves coriaceous, about 1 cm broad, falcate; styles with 2-4 stigmatic arms;

anthers attached at equal heights 1. Cymodocea

Leaves grass-like, linear to setaceous; style simple, attached laterally to the globose

carpel; one anther situated above the other 2. Halodule

Brackish or fresh water plants; monoecious; pollen globose:

Anther 2-4 locular; leaves opposite or alternate, linear; fruits curved, pectinate, apex

caudate 3. Zannichellia

Anther 1-locular; leaves crowded above, setaceous; fruits straight, smooth, with a

long pin-like apical appendage 4. Althenia

60 1. CYMODOCEA

Cymodocea Koenig in Koen. & Sims, Ann. Bot. 2 : 96, t.7 (1805); Aschers. in Pflanzenfam.

2, 1 : 210 (1889); A. Benn. in F.T.A. 8 : 228 (1901); Graebn. in Pflanzenr. 4, 2 : 146 (1907);

MacnaeinNat. Hist, oflnhaca Island, Mozambique 28, fig. 8f ( 1 958). Type species: C. nodosa

(Ucria) Aschers.

Submerged marine perennials with long, creeping, jointed, woody or fibrous, bracteate

rhizomes. Roots 1-6, arranged in a regular pattern which is typical for each species, branched,

densely covered with semi-persistent roothair-bases on the younger parts. Rhizomes sympodial

or simple. Stems erect, short or long, covered with prominent transverse leafscars. Leaves in

apical, distichous clusters, early deciduous, lamina strap-shaped or ribbon-like, straight or

falcate, coriaceous or thin, stipular sheath adnate to the lamina, amplexicaul at the base,

folded. Flowers dioecious, on much abbreviated lateral branches just below the leaf clusters,

surrounded by reduced leaves (bracts). Male flowers consisting of 2 fused stamens with the

2 pairs of apiculate locules facing opposite directions (filaments presumably lengthening

rapidly during anthesis and soon disintegrating as in Zannichellia); pollen filiform. Female

74

Zannichelliaceae

flowers consisting of 2 sessile, ovoid, longitudinally compressed, collateral carpels, with a short

style and 2 long subulate stigmatic branches, ovule solitary, pendulous. Fruit not well known.

Seven species are found in pantropical waters; one occurs in America, the others are Old World species.

One species is recorded from Zululand.

Cymodocea ciliata ( Forsk .) Ehrenb. ex

Aschers. in Sitzb. Ges. Naturf. Fr. Berlin 3

(1867); Linnaea 35 : 162 (1867), pro parte;

Schweinf., Beitr. FI. Aethiop. 292, 309 (1867);

Aschers. in Pflanzenfam. 2 : 210, fig. 163

(1889); Dur. & Schinz, Consp. FI. Afr. 5 ; 500

(1895); A. Benn. in F.T.A. 8- : 229 (1902);

Graebn. in Pflanzenr. 4, 2 : 151, fig. 33, A,

B (1907); Macnae in Nat. Hist, of Inhaca

Island, Mozambique 28 fig. 8f. (1958); Ward

in Lammergeier 2 : 21 (1962). Type: Egypt,

Red Sea, Forskal (CP. BM).

Zostera ciliata Forsk., FI. Aeg.-Arab. 157 (1775).

Thalassia cil ata (Forsk.) Koenig in Koen. &

Sims, Ann. Bot. 2 : 97 (1806); Kunth, Enum.

3 : 120 (1843).

Phucagrostis ciliata (Forsk.) Ehrenb. & Hempr.,

Symb. Phys. Bot. t.6, ined., ex Aschers. l.c.

Perennial, woody, rhizomatous herbs,

glandular-punctate. Roots 6 (occasionally

some bifurcating) appearing in a circle from

definite points on the fourth node and inter-

node of each section of the sympodial rhizome,

long, much-branched, forming dense mats,

woody, with long scattered root-hairs their

thickened bases persisting as short bristles.

Rhizome sympodial, each section consisting of

4 intemodes, the fourth forming the erect

stem, a lower bud continuing the rhizome;

internodes angled, smooth, orange brown,

with deltoid, amplexicaul, early deciduous

bracts. Stems erect, up to 50 cm, unbranched,

or rarely with a short side branch, with pro-

minent annular, curved leaf-scars, the lowest

internodes up to 15 mm long, the upper

shorter, 1-2 mm long, flattened, orange-

brown, occasionally forming a few rootlets

below. Leaves about 6-10, in distichous apical

tufts, early deciduous, strap-shaped, falcate,

7-15 cm long and 7-15 mm broad, leathery,

margin with erect teeth in upper half and on

the obtuse to truncate apex, midrib distinct

and with about 10 lateral nerves on each side;

below the lamina is attached to a light brown,

punctate, flat, stipular sheath which is tapered

to the base, with folded margins and a narrow

rim-like ligule above; axillary scales seta-

ceous, 3-6. Flowers on much abbreviated side

branches near the base of the leaf-clusters;

the leaves also reduced in size to form bracts.

Male flowers consisting of 2 fused stamens

with the 2 pairs of apiculate locules facing

opposite directions; filaments (or pedicels)

presumably lengthening rapidly during an-

thesis and soon disintegrating, pollen fila-

mentous. Female flowers at first sessile,

peduncle accrescent, knobbed, 4-14 mm

long, flattened, erect, adjacent to the main

stem with 2 fertile pairs of bracts, the inner of

a pair about ^ of the outer in size; the 2 col-

lateral carpels situated on the apex of the

peduncle, surrounded by groups of setae

(axillary scales), carpels ovoid, compressed

lengthwise with a short style 4 mm long

divided into 2 subulate stigmatic arms about

25 mm long, ovule solitary, pendulous. Fruit

(immature?) cylindrical, about 15 mm long,

2 mm broad, apex truncate. Fig. 21 : 1.

Distributed from the Red Sea eastwards in the

Indian Ocean to the tropical Asiatic coasts of the

Pacific and the tropical east coast of Australia;

southwards along the tropical and subtropical

east coast of Africa as far as Zululand. Common

and plentiful to dominant on the tropical east

African coast, in the infratidal zone, adhering to

rocks or coral reefs, rarely uncovered for a short

period at very low tides. The colonies are usually

unisexual, rarely bisexual. In Zululand it is found

attached to sandstone rock-pools exposed to wave

action.

Natal. — Ingwavuma: Mabibi Rocks, 33 miles

S. of Mozambique border, Tinley & Ward 24;

Kosi Estuary, Edwards 3611; Black Rock, Ward

3710; sandstone reef near Big Kosi Lake, Rodin

4627.- Ubombo: Sordwana Bay, Jesser Point,

Ward 3357. Lower Umfolozi: Mapalane, Ward

3929; 3930; 3391.

There is a marked difference in the appearance

of plants growing on sheltered warm coral reefs,

where there is no wave action, and those found in

sandstone rock crevices and in pools on beaches

exposed to waves and to lower temperatures, which

happens to be the habitat of the South African

plants. These plants show gnarled rhizomes, more

flexible stems and thinner narrower leaves; more-

over the orange colour is absent. The description

given here is based on the more typical form from

the sheltered, warmer coral reefs. Anatomically

the various parts of the two forms showed very

little difference. There were 8-10 vascular bundles

in the typical form, whereas the more slender form

developed only 6-8 bundles in the cortex of the

stem and rhizome. The internodes of the stem are

Zannichelliaceae

75

Fig. 21. — 1, Cymodocea ciliata, x 1; la, leaf-tip, x 5. 2, Althenia filiformis, X 1; 2a, female and male

flowers, X 6; 2b, unilocular anther, x 12; 2c, drupe, x 10.

76

Zannichelliaceae

longest below but shorten upon reaching water

level. Sheltered plants become taller than those

exposed to much wave action.

A small structure appearing to be a young fruit

was seen on Moss 20674 from Inhaca, Mozambique.

It was oblong, flattened, 15x2 mm, truncate and

apiculate above, ribbed (perhaps through desic-

cation) and glandular.

So far all collections from the Republic have

been sterile, possibly because of the water being too

cold. Ward, who studied the species in its natural

habitat, suspects that it does not tolerate being

covered by sand. As a result it generally occurs at

the northern and north-eastern sections of the sand-

stone outcrops in these bays. The greatest depth

recorded by him was 40 cm below the lowest

water level in open habitats, 64 cm in a pooh

61 2. HALODULE

Halodule End!., Gen. Suppl. 1 : 1368 (1841); Aschers. in Linnaea 35 : 163, 187 (1868);

Pflanzenfam. 2, 1 : 212 (1889); Sauvageau in Journ. Bot. France 4 : 321 (1890); den Hartog

in Blumea 12 : 289 (1964). Type species: H. uninervis (Forsk.) Aschers.

Diplanthera Thouars, Gen. Nov. Madag. 2 : 3 (1806); Steinheil in Ann. Sci. Nat. 2, 9 : 98 (1838); Aschers.

in Pflanzenfam. Nachtr. 37 (1889); Graebn. in Pflanzenr. 4, 2: 151 (1907); non Gleditsch (1764).

Halodula Benth. & Hook.f., Gen. PI. 3 : 1019 (1883).

Perennial slender grass-like herbs with a creeping rhizome, rooting at the nodes. Leaves

distichous, 2-4 per shoot, linear to setaceous, straight, flat, dark green, midrib and margins

ending in short apical teeth; stipular sheath long, attached to the lamina, convolute, flat,

forming 2 rounded auricles above, ligule a narrow apical ridge. Flowers dioecious, unisexual,

solitary, on a short side branch, surrounded by membranous leaf-sheaths. Male flowers

consisting of a double fused stamen with the bi-locular, sometimes apiculate, anthers inserted

at different heights facing opposite directions, attached dorsally to the filament, dehiscing

longitudinally, occasionally with 3 small protuberances inserted at different heights on the

connective, pollen confervoid. Female flower consisting of 2 collateral carpels on a short

clavate verrucose peduncle, style long, subulate, ovule solitary, pendulous. Fruit a globose,

hard berry.

A pantropical marine genus consisting of 7 closely related species usually forming extensive sea-meadows

on intertidal sandbanks. One species in South Africa.

Halodule uninervis (Forsk.) Aschers. in

Boiss., FI. Orient. 5 : 24 (1882); Pflanzenfam.

2. 1 : 213 (1889); Sauvageau in J. Bot. France

4 : 321 (1890); den Hartog in Blumea 12 : 297

(1964). Type: Yemen, near Mocha Coast,

Forskal (type could not be located).

Zostera uninervis Forsk., FI. Aeg.-Arab. CXX>

157 (1775).

Diplanthera uninervis (Forsk.) Aschers. in Pflanzen-

fam. Nachtr. 37 (1897); Graebn. in Pflanzenr.

4, 2 : 152 (1907); Macnae in Nat. Hist, of Inhaca

Island, Mozambique 28, fig. 8a (1958).

Cymodocea australis sensu A. Benn. in F.T.A.

8 : 229 (1902).

Grass-like perennials. Roots up to 6,

arranged in a semi-circle at the nodes of the

main rhizome, simple, long, densely covered

with long root-hairs; the roots from the

abbreviated nodes of the rhizome-branches

appearing singly, inserted left and right to

form a double row. Rhizome with internodes

up to 4 cm long, giving off much abbreviated

side branches with internodes 0 • 5-3 mm long,

ending in a leafy shoot; the internodes

covered at first with membranous long, im-

bricate, amplexicaul, convolute bracts, their

blackish bases persistent, giving the rhizome-

branches their typical beaded appearance.

Leaves 2-4 on very short, erect shoots; leaf-

sheaths (the lower without a lamina) long,

folded, bi-auriculate; lamina linear to seta-

ceous, straight or somewhat falcate, flat,

dark green, c.100 mm long and 2 mm broad,

midrib and margins terminating above in 3

short teeth, the truncate area between the

teeth sometimes slightly convex or concave,

often with a dark spot below the median

tooth; secondary nerves indistinct, lamina

with scattered tannin cells. Flowers solitary,

beside a young leaf, surrounded by mem-

branous, somewhat dilated sheaths; male

pedunculate with one anther inserted above

the other, the locules facing opposite direc-

tions; female with 2 carpels situated collater-

ally on a clavate peduncle which bears 3 small

apical protuberances; style inserted laterally.

Zannichelliaceae

77

long, subulate, ovule solitary, pendulous.

Fruit globose, 1 • 5 mm in diam., hard, orange,

papillate. Fig. 22 : 1 .

Tropical east Africa from the Red Sea to Zululand,

Madagascar, Asia. Recorded once from the Republic.

Natal. — Lower Umfolozi: Mapelane area, sand-

stone shelf in upper infratidal zone, Ward 3931

(sterile).

The drawings of the flowers were made from

fertile material collected on Inhaca Island in

Mozambique. The male flower belongs to Mogg

27172, the female to Mauve & Verdoorn 23. Both

specimens agree with den Hartog’s concept of

Halodule wrightii, being finer and smaller plants.

It is likely that the peduncle or filament of the male

flower will elongate at anthesis. The stigmas of

the female flower had degenerated and could not

be observed.

62 3. ZANNICHELLIA

Zannichellia L., Sp. PI. 969 (1753); Gen. PI. ed. 5 : 416 (1754); Aschers. in Pflanzenfam. 2,

1 : 213 (1889); A. Benn. in F.C. 7 : 50 (1897); F.T.A. 8 : 224(1901); Graebn. in Pflanzenr-

4,2:153(1907); Adamson in FI. Cape Penins. 35 (1950); Phill., Gen. ed. 2 : 54(1951); Rendle,

Class. Flow. PI. ed. 2, 1 : 208 (1953); Hutch., Fam. Flow. PI. ed. 2, 2 : 559 (1959); Clapham,

Tutin & Warburg, FI. Brit. Isles ed. 2 : 960 (1962); Singh in Bot. Gaz. 126 (2) : 141 (1965).

Type species: Z. palustris L.

Submerged annuals, perennating occasionally. Roots 1-3 per node, simple, densely

covered with long root-hairs, penetrating the substratum spirally. Rhizome creeping, matted

sympodial, giving off simple or sparsely branched erect stems. Leaves ternate, opposite or

alternate, linear, sessile, with a free cupular sheath subtending the axils, or the leaf base

expanded into a folded sheath forming auriculate stipules above; axillary scales 2, minute.

Flowers monoecious, unisexual solitary, or with a collateral vegetative bud or one of the

opposite sex beside it; males ephemeral, on long filaments (pedicels) anthers 4-8-locular,

with an apical knob; pollen globose, copious; female surrounded by a spathe, solitary or

pseudo-umbellate, ovary surrounded by a closely appressed flask-shaped “perianth”, style

exserted, tubular, stigma spoon- or funnel-shaped, large, with wavy membranous margins,

ephemeral, the ovule pendulous. Fruit a somewhat bean-shaped drupelet, stipitate and

crowned by the persistent style, pectinate on the dorsal and occasionally on the ventral side;

seed with a long, spirally coiled hypocotyl, exendospermous.

Two species; one a widespread variable species (regarded by some as multispecific) the other endemic

to the Cape Peninsula and neighbouring coastal areas. They inhabit brackish and fresh water and multiply

rapidly.

Flowers umbellate; leaves usually ternate or opposite, sheaths cupular, free from lamina (section

Zannichellia) 1 . Z. palustris

Flowers solitary; leaves alternate, lamina expanded below to form a convolute sheath (section Pseudal-

thenia) 2. Z. aschersoniana

1. Zannichellia palustris L., Sp. PI. 969

(1753); Aschers. in Pflanzenfam. 2, 1 : 213

(1889); A. Benn. in F.C. 7: 50 (1897); Camp-

bell in Proc. Calif. Acad. Sci. ser. 3, Bot.

1 (1897); Graebn. in Pflanzenr. 4, 2 : 153

(1907); Adamson in FI. Cape Penins. 35

(1950); Rendle, Class. Flow. PI. ed 2, 1 : 208,

fig. 83 c-e (1953); Hutch., Fam. Flow. PI. ed.

2, 2 ; 559, fig. 355 (1959). Type: Europe,

(LINN 1085.1).

Z. palustris subsp. pedicellata Wahlenb. & Rosen

in Nov. Act. Upsal. 8 : 227, 254 (1821); Graebn.

in Pflanzenr. 4, 2 : 156 (1907); Marloth in FI.

S. Afr. 4, 10, fig. 2 (1915). Z. pedicellata Fries,

Novit, Mant, 3 : 133 (1842); Wager in Trans.

Roy. Soc. S. Afr. 16 : 202 (1928). Type: Europe-

Z. stylaris Presl, Bot. Bemerk. 112 (1844). Type:

Cape, Drege 8801. Z. palustris var. pedicellata

Aschers. in FI. Brandenb. 669 (1864); Dur. &

Schinz, Consp. FI. Afr. 5 : 498 (1895).

Submerged brittle annuals forming dense

mats resembling fine grass. Rhizome thin,

branching freely. Stems erect, thin, up to

50 cm long but usually shorter. Leaves 1-3

per node, linear, 2-5 cm long and 1-2 mm

broad, gradually tapered above, below sur-

rounded by a free tubular, membranous,

scarious sheath; axillary scales minute,

lanceolate. Flowers monoecious, unisexual,

78

Zannichelliaceae

Fig. 22. — 1, Halodule uninervis, x 1; la, leaf-sheaths, X 2; lb, lc, leaf-tips, x 4; Id, young male flower,

x 9; le, female flower, x 10. 2, Zannichellia palustris, x 1; 2a, diagrammatic drawing of immature

flowers, X 10; 2b, 4-locular stamen showing small pustule, x 22; 2c, fruit, X 8.

Zannichelliaceae

79

the male and female flowers aggregated in the

axils of the leaf whorls, occasionally the

upper axils producing only males, each sur-

rounded by a membranous tubular sheath.

Male flower consisting of a solitary stamen;

filament lengthening rapidly at anthesis, up to

2 cm long, anther 4-8 locular, with a glandu-

lar apical knob and a small pustule at the base

of the connective on the axial side, pollen

globular. Female flowers 1-6, arranged in a

pseudo-umbel, enclosed at first in a cupular

spathe; peduncle 1-5 mm long, carpel ovoid,

stipitate, style tubular, stigma spoon-shaped,

soon disintegrating. Drupelet bean-shaped,

2 mm, crested and knobbed on the convex

side, (occasionally also ventrally) apex pin-

shaped (the style and stigma-base being per-

sistent), stipitate. Fig. 22 : 2.

Cosmopolitan but absent from Australia. Wide-

spread but not common in Southern Africa; occurs

in fresh or brackish water.

Cape. — Bathurst: Kowie, Britten 5254. Coles-

berg: Stormsfontein, near Colesberg, Zeyher 1888.

Cradock: Sister Mary Francis. Hay: Niekerkshoop,

Marloth 4912. Herbert: Campbell, Wilman sub

KMG 1393. Kimberley: Pretorius Vlei, Hafstrom

1070. Middelburg: Grootfontein College, Comins

843. Oudtshoorn: near Kamenassie, Britten 3161.

Port Elizabeth: Swartkops River, Sc/ilechter 6080.

Williston : Walkraal dam, Foley 201.

O.F.S. — Bloemfontein: Moran. Fauresmith:

Fauresmith Reserve, Verdoorn 1574.

Transvaal. — Brakpan: Wager.

S.W.A. — Grootfontein: Abenab, Dinter 7403;

Namutoni, De Winter 2981. Maltahohe: Nomtsas,

Giess 2283. Okahandja: Gross Barmen, Wiss &

Kinges 998. Swakopmund: Nonidas, Swakop River,

Gal pin & Pearson 7447.

Campbell (l.c. 1897) gives a detailed account of

the morphology of Zannichellia palustris, especially

of the development of the flower. He found that

the stem-apex divides dichotomously, the one half,

forming the floral part, being recognizable by its

broader shape. This is due to a second dichotomous

division giving rise to a second flower. This splitting

may be repeated three or four times and as a result

2-8 flowers may appear. The spathe, or cup-like

envelope at the base of the flower cluster, should be

considered the homologue of the bracts found on

the nodes of the main axis. It is also believed to be

the equivalent of the Aroid spathe.

When examining the root, the presence of a

definite dermatogen layer extending over the root-

cap was observed by Campbell. This is not very

common. The axillary scales (squamae intravagi-

nales) were thought to be secretory organs, giving

off the mucilage so often encountered in the younger

parts of the plants.

2. Zannichellia aschersoniana Graebn. in

Pflanzenr. 4, 2 : 157 (1907); Adamson in FI.

Cape Penins. 35 (1950); Reynecke in Journ.

S. Afr. Bot. 30 : 93 (1964). Syntypes: Cape

Peninsula, Wollev Dod 3586; 3381 (B|, PRE,

BOL).

Annuals with slender rhizomes. Sterns

upright, up to 60 cm long, 0-75 mm in diam.,

usually unbranched. Leaves alternate, linear,

2-7-5 cm long and 1-2 mm broad, narrowed

gradually to the rounded apex, expanded

below to form a sheath 5 mm long, which is

auriculate above. Male flower a single stamen

at first enveloped in a thin, membranous

spathe, filament lengthening rapidly at an-

thesis, up to 25 mm long, the 4 locules forming

a rectangular anther (squared in cross-

section), dehiscing longitudinally, with a

rounded apical knob and 2 small basal

pustules on opposite sides of the connective.

Female flowers surrounded by a vaginate

spathe and an inner flask-shaped “perianth”

which is truncate above and closely appressed

to the ovary and immature style; during

anthesis the style lengthens rapidly and dis-

plays a large, ephemeral spoon- or funnel-

shaped stigma. Fruit an asymmetrically

graniform to somewhat bean-shaped drupelet,

c.4 mm long, dorsally and occasionally

ventrally, pectinate, stipitate below, crowned

by the persistent style-base, brownish orange.

The fruits formed below on the rhizomes

geotropic, borne on stalks usually about 2 cm

long (up to 6 cm) ; the fruits at right angles to

the stalk. Fig. 23.

Endemic to the Cape Peninsula and neighbouring

coastal areas; northwards it is reported from

Famberts Bay, eastwards from Still Bay; in

brackish pools near the sea.

Cape. — Caledon : near Gordons Bay, Parker 4923.

Clanwilliam: Famberts Bay, Tange Vlei, Tolken

333. Peninsula: Kommetjie, Wolley Dod 3381;

Noordhoek Flats, Mauve & Simons 4251; Rietvlei,

near Milnerton, Stephens (51 of waterplant series);

Mauve & Simons 4252.

63 4. ALTHENIA

Althenia Petit in Ann. Sci. Obs. 1 : 451 (1829); Aschers. in Pflanzenfam. 2 : 213 (1889);

Aschers. & Graebn., Synops. Mitteleurop. FI. 1 : 365 (1897); Graebn. in Pflanzenr. 4, 2 : 158

(1907); Schonland in Kew Bull. 365 (1924); Adamson in FI. Cape Penins. 36 (1950); Phill .,

Gen. ed. 2 : 54 (1951); Hutch., Fam. Flow. PI. ed. 2, 2 : 559 (1959).

80 Zannichelliaceae

Fig. 23. — 1, Zannicheliia aschersoniana, X 1; a, young male flowers, x 6; b, cross-section of anther;

c, young female flower, x 10; d, older female flower; e, fruit, x 6.

Zannichelliaceae

81

Very small, creeping, submerged annuals. Roots 4-7 from a node, simple, long, densely

covered by long root-hairs. Rhizome branched, slender, with a deciduous, lanceolate, am-

plexicaul scale from each node. Stems simple, erect, up to 15 cm long, filiform. Leaves usually

aggregated above, filiform-setaceous, up to 5 cm long, sheaths folded, membranous, about

5 mm long, above forming a forked appendage 2-3 mm long, becoming frayed with age.

Flowers aggregated above. Male flowers solitary, on a pedicel which elongates up to 4-5 mm

during anthesis so that the anther is just exserted; often with 3 as yet undeveloped female

flowers beside them; perianth a small 3-toothed cup, with the unilocular anther sessile or

subsessile, pollen globose. Female flowers usually 3 together, ovary stipitate, enfolded in a

membranous bract, narrow-ovoid, unilocular, with a long style and a saucer-shaped apical

stigma. Fruit a stipitate, cylindrical, slightly oblique, drupelet crowned by the persistent

pin-like style and stigma-base.

A monotypic genus found in southern Europe, northern and southern Africa.

Althenia filiformis Petit in Ann. Sci. Obs.

1 : 451 (1829), etc. l.c. Type: France.

Description as for the genus. Fig. 21 : 2.

Found in brackish pools near the sea on the

western and southern coasts of Southern Africa.

Cape.- — Humansdorp: mouth of Kabeljous River,

Gillett 2334. Namaqualand: Hondeklip Bay,

Pillans in BOL 18258. Peninsula: Paarden Eiland,

Adamson 3391 ; Rugby, Tolken 445. Port Elizabeth :

Chatty River near Redhouse, Holland sub Schon-

land 4450; Veeplaats, saltpan, Archibald 5018.

S.W.A. — Liideritz: Liideritzfelder Teiche, Dinter

6414.

It was observed that the bracts surrounding the

flowers are modified leaf-sheaths, the ligules

especially being well developed. All stages between

the normal leaf and the bract with a tiny caudate

appendage, representing the lamina, were observed.

Schonland in his note on the discovery of this

species in South Africa (Kew Bull. 1924, p.365)

failed to detect the unilocular stamens, but they are

present in the Holland collection from the Chatty

River.

NAJADACEAE

by A. A. Obermeyer

Submerged herbaceous annuals or perennials, with simple adventitious roots devoid of

root-caps. Upright branches forked repeatedly, brittle, smooth or prickly. Leaves sessile,

in psuedo-whorls of 3 or more; with an open folded basal sheath and a linear dentate lamina;

with 2 axillary scales. Flowers monoecious or dioecious, solitary or sometimes aggregated

above, unisexual, small. Male flower a single stamen enveloped in bud by a closed spathe;

closely adhering to the 1-2 or 4-locular anther is a thin inner envelope which is bilobed at

the apex; pollination hydrophyllous, pollen globose or ellipsoid, without exine, contents

granular. Female flower a single ovoid carpel surrounded by a spathe or naked; ovule solitary,

basal, erect, anatropous, style filiform, with 2-3 stigmatic branches (viz. in Old World species).

Fruit a small narrow-ovoid nut, epicarp reticulated, embryo straight, with a large hypocotyl

and radicle, exendospermous.

Monogeneric.

82

Najadaceae

64 NAJAS

Najas L., Sp. PI. 1015 (1753); Gen. PI. ed. 5 : 445 (1754) ; A. Braun in Journ. Bot. Lond. 2 : 274

(1864); Magnus, Beitr. z. Kennt. Gatt. Najas (1870); Pflanzenfam. 2, 1 : 214-218 (1889);

Benth. & Hook. f. Gen. PI. 3 : 1018 (1883); Rendle in Trans. Linn. Soc. Bot. 2, 5 : 379-444,

t. 3 9-42 (1899). A. Benn. in F.C. 7 : 51 (1897); F.T.A. 8 : 225 (1902); Rendle in Pflanzenr. 4,

12 : 1 (1901); Phill., Gen. ed. 2 : 55 (1951); Lawrence, Taxonomy of Vascular Plants 378

(1951); Hutch., Fam. Flow. PI. ed. 2, 2 : 561 (1959); de Wilde in Acta Bot. Neerl. 10 : 164-170

(1961); FI. Males, ser. 1, 6 : 157 (1962). Wild in Kirkia 2 : 37 (1961). Type species: N.

marina L.

Description as for the family.

About 35 species widely distributed but absent from very cold areas. Found in fresh or brackish water;

3 widespread species in Southern Africa.

Several species of this genus were closely studied by botanists in Europe and in America, e.g. Magnus,

Schumann, Rendle, Bailey, Campbell and recently, de Wilde. Campbell noted the absence of a root-cap

in the American species, N. flexilis. In N. pectinata I found no root-hairs. The “ternate” phyllotaxis is

made up of two subopposite amplexicaul leaves while the third is one of a “pair” produced in the axil of

the lower leaf: the other leaf of this “pair,” however, does not develop but remains minute and scale-like;

in its axil is found a vegetative bud or a male or female flower.

De Wilde, in his studies of Asian material, found that on occasion the male or female flower developed

a spathe, although they were normally naked in these species. Furthermore he observed the different stages

in the transformation of an open scale to a closed, bottle-shaped spathe, concluding that it must be consi-

dered the homologue of a leaf.

Campbell noticed how the male and female flowers of A. flexilis were strikingly similar in the early stages,

both developing from an apical bud which divides dichotomously to form a flower and a sterile branch.

Besides the outer envelope, or spathe discussed above, the young stamen develops an inner envelope which,

according to Campbell, might be the homologue of the ovular integument. Magnus and Rendle on the

other hand interpret this inner envelope as a perianth.

Stems usually with scattered prickles, succulent; septa of leaf indistinct; stigmas 3; usually found in

brackish water (Section Najas) 1 . N. marina subsp. delilei

Stems smooth, herbaceous; septa of leaf distinct; stigmas 2; found in fresh water (Section Caulinia):

Leaves with about 6-12 macroscopic triangular, spinetipped teeth; dioecious; male flower sur-

rounded by a sheath 2. N. pectinata

Leaves filiform with 40-60 microscopic teeth; monoecious; male flower naked 3. N. graminea

1. Najas marina L„ Sp. PI. 1015 (1753).

Type: Europe (LINN 1 156.1-2).

subsp. delilei ( Rouy ) Maire in FI. Afr.

Nord 1 : 205 (1952). Type: Lower Egypt,

Fareskour, Delile (MPU, holo.; PRE,

photo.).

N. delilei Rouy in FI. France 13 : 294 (1912),

norm. nov. N. muricata Del., FI. Egypte 281, t.50.

fig. 1 (1813), non Thuillier (1799). Type: as for

N. delilei. N. marina L. var. muricata (Del.) A.

Braun ex K. Schum. in FI. Bras. 3, 3 : 725 (1894);

Rendle in Trans. Linn. Soc. Bot. ser. 2, 5 : 397,

440 (1899); Pflanzenr. 4, 12 : 8 (1901); A. Benn.

in F.T.A. 8 : 226 (1901). N. armata Lindb. f. in

Acta Soc. Sci. Fenn., B, 1, 2 : 8 (1932), nom. nov.

for N. muricata Del.; Tackholm, FI. Egypt 1 : 113

(1941). N . marina subsp. armata (Lindb. f.) Horn af

Rantzien in Kew Bull. 1952 : 29 (1952).

Caulinia muricata (Del.) Spreng., Syst. 1 ; 20

(1825).

Annuals up to 60 cm high, somewhat

succulent, often decumbent in shallow water,

rooting at the nodes. Stems (in S.A. speci-

mens seen) c. 2 mm in diam., spinoso-dentate

or occasionally smooth, prickles similar to

those of the leaves; lower internodes up to

7 cm long, upper shorter. Leaves linear,

c. 20 mm long and 2 mm broad, the margin

and often also the midrib dorsally spinoso-

dentate with the apical cell of the spine sharp,

brown; basal sheath small, 4 mm. auriculate,

membranous, obtuse, denticulate or smooth.

Flowers dioecious, solitary; male surrounded

by a bottle-shaped spathe contracted into a

thin denticulate, oblique neck; stamen 2-4

celled, the septa often incompletely developed,

pollen ellipsoid; female naked, with an ovoid

ovary 3 mm, style with 3 erect stigmatic papil-

late branches about half as long as the ovary.

Najadaceae

83

Fruit an ovoid nut 4 mm long, coarsely honey-

combed. Fig. 24 : 2.

North-western Europe, Africa, Asia to western

Australia. Rare in Southern Africa; in brackish

pools in eastern coastal belt; also recorded from

the Kruger National Paik. Apparently selective

in its habitat. (N.B. Not a marine species).

Cape. — George: Lange Vlei, Stephens 30. Knysna:

Karatara Lake, Martin 4514.

Natal. — Ingwavuma: Nyamiti Lake, Ward 3208;

Nhlange Lake, Tinley 356.

Transvaal. — Letaba: Kruger National Park,

Tsende River, Brynard & Pienaar 4312.

2. Najas pectinata (Pari.) Magnus in

Illustr. FI. Egypte (Mem. Inst. Egypt. 2, 1)

145 (1889); Ber. Deutsche Bot. Ges. 12 : 219

(1894); Dur. & Schinz, Consp. FI. Afr. 5 : 500

(1898); Horn af Rantzien in Kew Bull.

1952 : 38 (1952); Wild in Kirkia 2 : 37 (1961).

Type: Egypt, Medinet-el-Fayoum, Figari

(FI, holo., PRE, photo.).

Caulinia pectinata Pari., LI. Ital. 3 : 665 (1858).

Najas horrida A. Braun ex Rendle in Trans.

Linn. Soc. ser. 2, 5 : 422, t.42, fig. 183-191 (1899);

ib. 443 (1900); Pflanzenr. 4, 12 : 17 (1901). Type:

Nigeria, Nupe, Barter 1065. N. interrupta K.

Schum. in Pilanzenw. Ost. Afr. C, 94 (1895);

A. Benn. in F.C. 7 : 51 (1897); F.T.A. 8 : 228

(1902); Rendle in Trans. Linn. Soc. ser. 2, 5 : 423

(1899); Pflanzenr. 4, 12 : 17 (1901); Marloth,

FI. S. Afr. 4 : 9, fig. la (1915); Horn af Rantzien

in Kew Bull. 1952 : 38 (1952). Type: Tanganyika,

Lake Victoria, Fischer 614 (Bf, K, lecto).

Submerged perennial plants forming

dense brownish clumps 30-100 cm high, some

of the upper leaf-tips just exserted above the

surface of the water. Stems much ramified,

rooting at the nodes; upper stems erect, thin,

terete, smooth, 0-5-1 mm in diam., lower

internodes up to 12 cm long, abbreviated

above, the shoots appearing bushy. Leaves

aggregated in upper axils, spreading to falcate

from above the sheath, narrowly linear,

2-3 cm long, 0-5-1 mm broad, spinoso-

dentate, with 6-18 yellow, sharp teeth which

curve upwards on each side, sometimes teeth

innocuous and small, apex attenuate, spinous,

septa distinct; sheath usually auriculate,

amplexicaul, minutely dentate above, 3 mm

long. Flowers dioecious. Male flowers at first

enveloped in a tight membranous bottle-

shaped sheath with a few minute teeth sur-

rounding the aperture; stamen solitary, 3 mm

long; during anthesis the filament lengthens

rapidly up to 3 mm and the stamen bursts out

of the spathe above; inner envelope swollen,

forming 2 small pyramidal gibbosities above;

anther 4-lobular the integuments splitting

downwards and curling up outwards, the

inner septa remain standing as a 2-winged

column; the whole stamen soon disinte-

grating after shedding its pollen; pollen glo-

bose. Female flowers without a spathe, 2 mm

long, ovary turbinate, contracted into a short

style and 2 filiform stigmas. Nut cylindrical,

2 • 5 mm long, apiculate, marked with quadrate

pits. Fig. 24 : 1 .

A common inland fresh-water species found in

Natal, Transvaal, South West Africa and further

north to Egypt; also in Madagascar. Avidly eaten

by fish. Common name: Sawgrass.

Natal. — Hlabisa: Hluhluwe Game Reserve,

Ward 2566; 2945. Ingwavuma: Ndumu Game

Reserve, Tinley 457. Lower Umfolosi: Richards

Bay, fresh water lake, Wager.

Transvaal. — Brits: Hartebeestpoort Dam,

Talbot. Potgietersrus : Mosdene, near Naboomspruit,

Mauve 4201. Pretoria: Rietvlei Dam, near Pretoria,

Repton 2002; Rust de Winter, Wouda 4; Pienaars

River Dam, Mauve & Vahrmeijer 4303. Waterberg:

near Nylstroom, Prosser. Without locality, McLea

in BOL 6383.

S.W.A. — Okavango Native Territory : Okavango

River, Niagana, Dinter 7264; lagoon at Kapako

Camp near Mupini Mission Station, De Winter &

Marais 503 1 .

Although the species was assumed to be monoe-

cious by previous workers, the plants examined in

the herbarium and in the wild state were found to

be dioecious. Winterbuds in the form of thickened

nodes and leafbases were observed. A surprising

degree of desiccation can be tolerated.

The plant described as N. interrupta K. Schum.,

which is more slender and less spinous, appears to

be a deep-water form.

In many aquatic plants, e.g. Najas, Lagarosiphon

etc., the upper internodes are retarded in their

development, giving the plants the characteristic

bushy appearance. This retardation allows for

easy adaptation to a rising water-level.

3. Najas graminea Del., FI. Egypte,

2 : 282, t. 50, fig. 3 (1813); A. Benn. in F.T.A.

8 : 226 (1901). Type: Egypt, Rosetta Delta,

Delile (MPU, holo.).

Small, unarmed plants with much

branched shoots, 10-60 cm long, often

plumose above because of the closely packed

leaves. Roots many, long, filiform, with a

dense covering of long root-hairs towards the

tip. Stems with terete, brittle internodes,

1 mm in diam. Leaves 16-25 mm long, 0-5-6

mm wide, teeth microscopic, about 40-55 on

84

Najadaceae

Fig. 24. — 1, Najas pectinata, x 1; la, leaf, x 3; lb, leaf-sheath, x 7; lc, young male flower enclosed in

tubular spathe, x 10; Id, ripe male flower, x 10; le, female flower, x 25; If, seed, x 12. 2, N.

marina, x 1; 2a, leaf-sheath, x 5. 3, N. graminea, x 1; 3a, leaf-sheath, X 12; 3b, leaf margin,

x 100; 3c, male flower, x 12; 3d, female flower, x 12.

Najadaceae

85

each side; sheaths with linear-setaceous

auricles about 1 • 5 mm long, bearing some

minute teeth. Flowers monoecious. Male

flowers solitary, in upper leaf axils, naked;

pedicel at anthesis 1 mm long, anther 1 mm

long, oblong, apically with 2 small obtuse

ridges. Female flowers usually in lower axils,

sessile, about 1 • 5 mm long, the ovary about

as long as the short style and the 2 long stig-

matic arms. Fruit oblong, 1-5 mm. Fig.

24 : 3.

Collected once in Southern Africa but widely

distributed elsewhere. Recorded from Egypt,

tropical East Africa, Asia, Australia; introduced in

Italy and Britain.

Transvaal. — Pretoria: Wallmansthal, 20 miles

E. of Pretoria on farm Haakdoringfontein, Mauve

4342.

APONOGETONACEAE

by A. A. Obermeyer

Fresh-water aquatics. Rhizome tuberous (in South African species) densely covered with

thin, dead and living roots. Leaves submerged, aerial or with a floating blade, linear to elliptic,

with the base attenuate, obtuse or cordate, petiolate, innermost lateral nerves placed close to

the midrib, others more widely spacrd, transverse nerves at right angles to laterals, close

together (lamina fenestrated in some species from Madagascar). Spikes exserted, simple or

forked (rarely 3-8 fid in some species from Madagascar), enveloped in bud by a hood-shaped

spathe which is usually early caducous (persistent in a few species outside Africa). Flowers

ebracteate, bisexual and protrandrous or female, arranged dorsally or around the axis;

perianth-segments (or bracts) 1-3, persistent or deciduous (viz. in A. stuhlmannii and some

closely related species) white, pink, red, mauve or yellow, folded inwards in the budstage,

spreading afterwards, turning green with age if persistent. Stamens 6-16, free, hypogynous,

anthers small, bilocular, with the locules facing opposite directions, basifixed; pollen ellipsoid

to globose, pollination entonrophilous. Gynoecium of 3-6 (-9 in apomicts) free, sessile,

carpels, turbinate, attenuated into a style; style curving outwards and forming a stigmatic

ridge on the ventral suture; ovules 2-8, basal, anatropous. Follicles membranous, opening

on the adaxial side, the outer skin soon disintegrating; seeds fusiform, in some species with

a soft spongy outer covering, without endosperm, embryo straight.

Monogeneric.

65 APONOGETON

Aponogeton L./., Suppl. 32 (1781); Benth. & Hook.f., Gen. PI. 3 : 1013 (1880); Engler in

Bot. Jahrb. 8 : 261 (1887); Pflanzenfam. 2, 1 : 218 (1889); A. Benn. in F.C. 7 : 42 (1897);

F.T.A. 8 : 216 (1901); Krause & Engler in Pflanzenr. 4, 13: 1 (1906); Phill., Gen. ed. 2 : 55

(1951); Hutch., Fam. Flow. PI. ed. 2, 2 : 552 (1959). Type species: Aponogeton monostachyos

L.f.

Description as for the family.

A genus of about 25 species, widespread in Africa, Asia and Australia. Five species recognized in Southern

Africa.

86

Aponogetonaceae

Spike bifid:

Spikes with the flowers dorsally arranged in 2 rows; perianth-segments 1-2, spreading, longer than

the stamens; winter rainfall region:

Perianth-segment 1, large; spikes many-flowered, fragrant 1. A. distachyos

Perianth-segments 2; spikes 4-8-flowered, scentless 2. A. angustifolius

Spikes with the flowers arranged all around the axis or, if secund, not in 2 rows; perianth-segments

usually 3, equalling the stamens in length; summer rainfall region:

Lamina awl-shaped to lanceolate, base attenuated; scape not swollen above; flowers white,

pinkish or lilac, in fairly short secund or cylindrical spikes, obtuse at the apex 3. A. junceus

Lamina oblong-lanceolate, base cordate; scape swollen above; flowers yellow or white, in long

cylindrical tapered spikes 4. A. desertorum

Spike simple, few-flowered; perianth-segments deciduous, small; delicate fugitive plants

5. A. stuhlmannii

1. Aponogeton distachyos L.f., Suppl. 215

(1781); Thunb., Nov. Gen. 4 : 74 (1781);

Ait., Hort. Kew. 1 : 495 (1789); Andr., Bot.

Rep. 5 : t.290 (1803); Gawl. in Bot. Mag.

1. 1 293 (1810); Spreng., Syst. 2 : 465 (1825);

Engl, in Bot. Jahrb. 8 : 272 (1887); Dur. &

Schinz, Consp. FI. Afr. 5 : 492 (1894); A.

Benn. in F.C. 7 : 43 (1897); Krause & Engl,

in Pflanzenr. 4, 13 : 21 (1906); Marloth, FI.

S. Afr. 4 : 13, 14, tab.3, fig. 3 (1915); Ver-

doorn in S.A. Journ. Nat. Hist. 3, 2 ; 17

(1922); Adamson in FI. Cape Penins. 37

(1950). Type: Cape, T/mnberg (LINN 479 • 3,

pro parte, viz, the leaves and the spike second

from right, lecto.).

Tubers up to 6 cm in diam., dark. Leaves

with a floating lamina, lanceolate to ob-

long, 6-20 cm long and 2-7 cm broad, apex

acute to obtuse, base rounded or shortly at-

tenuate. Spikes bifid, many-flowered, up to

6 cm long. Flowers many, scented, dorsally

arranged in 2 spreading rows, with the flowers

on the one side slightly above those of the

other; perianth-segments solitary, oblong to

ovate-lanceolate, up to 18 mm long, delicate,

white or occasionally pink, accrescent and

green in fruit. Stamens 8-16, arranged in a

circle, filaments thin, 3 mm long, anthers

purple to black. Carpels 2-6 with c.4 ovules

in each carpel. Follicles turbinate, c.l cm long

with a straight beak, 1-4 seeded, seeds

9 mm long, with a spongy outer covering.

Fig. 25 : 2.

Common in the winter-rainfall area of the Cape

Province and, eastwards, to Plettenberg Bay; in

ponds, rivers and ditches. Often cultivated in Europe

and said to have gone wild in the south of France.

Flowers from March to September.

Cape.— Caledon : Elgin, Palmiet River, Smith

2554. Ceres: Verkeerde Vlei, Compton 12055.

Clanwilliam: Olifants River near Keerom, Pillans

8777. George: Diep River, Galpin 4778. Humans-

dorp: Fangkloof, Nuweplaas, Theron 1385; Ratels-

bosch, Fourcade 363. Knysna: Plettenberg Bay,

Rodin 1173. Laingsburg: Cabidu Dam, Compton

22892. Peninsula: Marloth 172; Rondebosch,

Schlechter 832. Riversdale: Vet River near Rivers-

dale, Muir 193. Stellenbosch: Faure, Strey 606.

Swellendam: Thode A2401.

Common names: Wateruintjie, Cape Hawthorn.

Both tubers and fruiting spikes are eaten. A culti-

vated plant of unknown origin (Schlieben 10499)

regularly produced simple spikes.

Dandy and Stearn, who investigated the typifi-

cation of A. distachyos , regard sheet 479.3 in the

Linnaean herbarium as the lectotype (although

comprising 2 species) for Linnaeus and his son

made annotations on the back of this sheet and

quoted Thunberg as the collector. The 3 spikes

on the left and the one on the right should be

excluded as they represent A. angustifolius. Although

sheet 479.2 consists of A. distachyos only, it cannot

be selected as the type since it was collected by

Sparrman.

2. Aponogeton angustifolius Ait., Hort.

Kew. 1 : 493 (1789); Willd., Sp. PI. 2 : 928

(1799); Gawl. in Bot. Mag. 1. 1 268 (1810);

Spreng., Syst. 2 : 465 (1825); Engl, in Bot.

Jahrb. 8 : 272 (1887); Dur. & Schinz, Consp.

FI. Afr. 5 : 492 (1895); A. Benn. in F.C. 7 : 43

(1897); Krause & Engl, in Pflanzenr. 4,

13 : 21 (1906); Marloth,>l. S. Afr. 4 : 13, 14,

t.3 (1915); Adamson in FI. Cape Penins. 37

(1950). Type: Cape Peninsula, Masson ,

introduced to Kew in 1788 (K, holo.).

A. crinifolium Lehm. ex Schlechtd. in Linnaea 10,

Litt. Ber. 76 (1836); A. Benn. in F.C. 7 : 45 (1897).

Type: Cape Flats, inundated places, Pappe (BM,

holo.; NBG).

Tubers globose, 2 cm in diam., brown.

Leaves: primary rarely developed, sub-

merged, filiform to linear, with a wavy margin,

tapered above and below; later leaves

petioled and with a floating lamina, narrowly

lanceolate to oblong, c.5 cm long and 1 cm

A PONOGETONACEAE

87

Fig. 25. — 1, Aponogeton angustifolius, X 1; la, flower. 2, A. distachyos, leaf a^d inflorescence, x 1; 2a,

flower, X 3; 2b, stamen, x 8; 2c, carpels, x 2.

4-F.S.A.

88

Aponogetonaceae

broad, apex acute, base rounded to attenuate,

lateral nerves indistinct, margin slightly

prominent. Spikes bifid, compact at first,

lengthening during anthesis, in fruit up to

2 cm long, laxly 4-8-flowered. Flowers scent-

less, situated dorsally on the rhachis;

perianth-segments 2, white with a pinkish

tinge below, oblong-lanceolate, 5-10 mm

long, accrescent in fruit. Stamens usually 6,

filaments subulate, 2-4 mm long, anthers

black. Carpels 2-4, ovules 2. Follicles narrow-

ovoid, green, membranous, 8 mm long, with a

straight beak; seeds 7 mm long, with a thin

spongy outer covering. Fig. 25 : 1.

Confined to the south-western Cape.

Cape. — Malmesbury: Darling, Bolus 12894;

Darling Flora Reserve, Winkler 98. Peninsula: near

Muizenberg, Schlechter 1253; Princess Vlei,

Walgate sub Adamson 3674; Grassy Park, Compton

13690; Isoetes Vlei, Barker 9748. Stellenbosch:

flats, Strey 602a, b. Worcester: Leipoldt s.n.;

Tolken 1072.

Linnaeus and his son confused this species with

A. distachyos. On sheet LINN 479.3 there are

4 flower spikes of A. angustifolius, but the leaves

and the second spike from the left belong to

A. distachyos (see p. 86).

3. Aponogeton junceusLe/tw. ex Schlechtd.

in Linnaea 10, Litt. Ber. 76 (1836); Steud.,

Nom. 114 (1841); “A. junceum Lehm.” Del.,

Sem. Hort. Hamb. (1883), nom. nud. “ A .

junceum herb. Zeyher” teste Hook.f. in Bot.

Mag. t.6399 (1878). Type: Hort. Hamburg,

ex Caffraria, Ecklon s.n.

A. spathaceus E. Mey. ex Hook.f., Bot. Mag. sub

t.6399 (1878); Engl, in Bot. Jahrb. 8 : 272 (1887);

Dur. & Schinz, Consp. FI. Afr. 5 : 493 (1894);

A. Benn. in F.C. 7 : 44 (1897); Rendle in Cat. Afr.

PI. Welw. 2 : 94 (1899); A. Benn. in F.T.A. 8 : 216

(1901). Krause & Engl, in Pflanzenr. 4, 13 : 1

(1906); E. Mey. in Drege, Zwei Doc. 165 (1843),

nom. nud.; Linnaea 20 : 215 (1847), nom. nud.

Type: Cape, Albert, Drege. — var . junceus (Lehm.

ex Schlechtd.) Hook.f. in Bot. Mag. t.6399 ( 1878) ;

A. Benn. in F.C. 7 : 45 (1901). Type: Hort. Hamburg

ex Caffraria, Ecklon (Bt). A. natalense Oliv. in

Hook., Ic. PI. t. 1471a (1884); A. Benn. in F.C.

7 : 44 (1897). Syntypes: Natal, York, McKen (K);

between Karkloof and Umgeni, Rehmanni K; BOL).

A. rehmannii Oliv. in Hook., Ic. PI. t. 1471b (1884);

A. Benn. in F.C. 7 : 44 (1897); Oberm., Schweick. &

Verd. in Bothalia 3 : 225 (1937). Type: Transvaal

Bushveld, between Kleinsmit and Kameelpoort,

Rehmann 4835 (K, holo.). — var. hereroensis

(Schinz) Krause & Engl, in Pflanzenr. 4, 13 : 16

(1906). — var. major Glover in Ann. Bol. Herb.

1 : 106 (1915), nom. nud., based on S.W.A., Great

Karasberg — Naruda Slid, Pearson 8456; 3457.

A. hereroensis Schinz in Bull. Herb. Boiss. ser.

2, 1 : 764 (1901). Syntypes: S.W.A., ponds east

of Windhoek, Dinter 587 (Z); just north of Water-

berg, Dinter (Z).

T ubers globose, up to 3 cm in diam. , greyish

brown. Leaves heterophyllous; filiform to

subtrigonous, 4-25 cm long; or apically ex-

panded into an erect spathulate lamina; or

a floating, oblong blade up to 17 cm long and

4-5 cm broad is formed, with the base and

apex attenuate or rounded. Spikes bifid,

usually 2-4 cm long and 8 mm broad (up to

9 cm long and 2-5 cm broad), lengthening

during anthesis, many-flowered. Flowers

bisexual or female, (through suppression of

stamens), scentless, situated dorsally on the

rhachis or only so near the base, with the

upper flowers arranged all around the rhachis ;

perianth-segments 1, 2 or 3, oblong-lanceolate,

white, pink or mauve, with a shiny, soft

texture, becoming green and firm with age.

Stamens usually 6, slightly longer than the

perianth-segments, equalling more or less the

carpels; filaments swollen at the base,

tapered above; pollen ellipsoid. Carpels up

to 7 in lower flowers, usually 3 in upper, ovoid,

with a curved beak above, stigmatic ridge

dark, ovules 1-8, basal. Follicles green, firm,

somewhat transparent, 3-5 mm long; seeds

fusiform, 2 mm long, outer spongy covering

thin; usually 1 or 2 seeds ripening per follicle.

Fig. 26.

Widespread in Southern Africa, with the exception

of the winter-rainfall area, and extending to

Rhodesia and Angola. Common in fresh-water

ponds, marshes and rivers, growing on the bank,

or submerged with the lamina floating and the flower

spikes exserted.

Cape. — Albany: Slaaikraal, near Grahamstown,

Mauve & Wells 24. Gordonia: Obobogorap, 120

miles N.W. of Upington, Leistner 1780. Kentani:

Pegler 1869. Mafeking: Moshesh near Mosita,

Bruckner 412. Maraisburg: 10 miles N. of Hofmeyr,

Acocks 16335. Riversdale: Oakdale, Muir 2747.

Uitenhage: Zeyher 1734.

O.F.S. — Fouriesburg: Witzieshoek, Junod in

TRV 17497. Kroonstad : Pont 590; Acocks 20994.

Basutoland. — Tsekilo’s, Jacot-Guillarmod 7551.

Mokhotlong, Thabana Ntlenyana, Coetzee 572.

Natal. — Hlabisa: Hluhluwe Game Reserve,

Ward 2200; 2493. Lions River: near Nottingham

Road, Edwards 2675. Maritzburg: Schlechter 3300.

Umzinto: Dumisa, Rudatis 1583.

Transvaal. — Bloemhof: Christiana, “Kaffraria”,

Burtt Davy 12798. Bronkhorstspruit : in river near

town. Mauve 4261 ; Repton 448. Carolina: Roberts

in TRV 14873. Petersburg: Brak River, Breme-

kamp & Schweickerdt 38. Pretoria: Wallmansthal,

20 miles N.E. of Pretoria, Dyer, Verdoorn <6 Mauve

Aponogetonaceae

89

Fig. 26. — 1, Aponogeton junceus, typical plant, x 1; a, form with floating leaves (“rehmannii”), X 1;

b, form with broad erect leaves (“natalense”), x };c, $ flower, x 10; d, stamen, x 6; e, young carpel,

x 4; f, fruiting carpel, x 3.

90

Aponogetonaceae

4273; Mauve 4344. Soutpansberg : farm Eyem,

north of Blaauwberg, Obermeyer, Schweickerdt &

Verdoorn 84. Waterberg: Mosdene near Naboom-

spruit, Galpin M.374.

S.W.A. — -Rehoboth: Buellsport, Klein Aub, Strey

2512. Grootfontein: Otavi, Dinter 5261; Merx-

miiller 2138.

The “species” A. junceus, A. natalense and A.

rehmannii are here regarded as forms of one

variable, complex species. They are high polyploids

and very likely unstable. Approximately 118-126

chromosomes were counted and many appeared to

be linked. The plants are often apomicts, the

stamens changing into carpels.

4. Aponogeton desertorum Zeyh. ex

Spreng.f, Syst. 5, Suppl. 14 (1828); Wild in

Kirkia 2 ; 36 (1961). Type; Cape, Uitenhage,

Zeyher 191 (B).

A. kraussiamis Hochst. in Krauss in Flora 343

(1845); Krauss, Beitr. FI. Cap. & Natal, 172

(1846); A. Benn. in F.C. 7 : 44 (1897); F.T.A.

8 : 217 (1901). Type: Cape, Uitenhage, Krauss

1604 (Bf; K). A. leptostachyus E. Mey. ex Engl,

in Pflanzenr. 4, 13 : 12 (1906); Drege, Zwei Doc.

165 (1844), nom. nud.; Engl, in Bot. Jahrb. 8 : 270

(1887), nom. subnud. Type: Cape, Uitenhage,

between Zuurberg and Bruintjieshoogte, Drege

(B). A. holubii Oliv. in Hook., Ic. PI. t.1470 (1884);

Dur. & Schinz, Consp. FI. Afr. 5 : 492 (1894);

A. Benn. in F.C. 7 : 44 (1897); F.T.A. 8 : 217

(1901); Oberm., Schweick. & Verd. in Bothalia

3 : 225 (1937). Syntypes: Bechuanaland, Henry’s

Pan. Holub 1036-39 (K). A. dinteri Engl. & Krause

in Bot. Jahrb. 36 : 92 (1905). Syntypes: South

West Africa, Otjimbingue, Fischer 165; Dinter ,

cultivated at the Botanic Garden, Berlin (Bf).

A. eylesii Rendle in Journ. Bot. Lond. 70; 76

(1932). Type: Rhodesia, Salisbury, Rua River,

Eyles 852 (BM, holo.).

Tubers small, globose, 1-2 cm in diam.,

brown. Leaves 6-9 per plant, petioles flat-

tened, striate, lamina floating, oblong, 5-17

cm long and 2-6 cm broad, apex obtuse,

apiculate, base cordate, thinly coriaceous,

often with many immersed oil glands in the

epidermis (no submerged leaves seen). Spike

bifid, lengthening during anthesis, 3—12 cm

long, scape flattened, usually swollen above.

Flowers white (in eastern Cape) or yellow,

eventually somewhat laxly arranged all around

the rhachis, the thick fruiting carpels below

and the elongated young apical part giving

the spikes their typically tapered appearance;

perianth-segments 2, narrow-oblong to

narrow-obovate, 2 mm long. Stamens usually

6, filiform, somewhat flattened, 2 mm long,

spreading, anthers small, early deciduous,

pollen ellipsoid, copious. Carpels 5-3, ovoid,

2 mm long, with a terete style, with the stig-

matic ridge short, apical, glutinous, usually

dark; the 4-8 ovules usually visible through

the transparent walls of the carpels and

follicles; seeds cylindrical, 2 mm long, mostly

about 6 ripening in each follicle. Fig. 27 : 1.

Recorded from the eastern Cape, Transvaal,

Bechuanaland and South West Africa; also in

Angola and Rhodesia. Found in rock pools or

temporary river pools, developing extremely fast

under favourable conditions. The flowers were noted

as white in the eastern Cape, but are yellow further

north.

Cape. — Albany: Committees, Dyer 2183.

Alexandria: Coerney River Valley, Zuurberg,

Archibald 5247. Bedford: near Bedford, Acocks

20286. East London: Rattray 259. Komga: near

Komga, Flanagan 1094; Gonubi Drift between

Komga and Kei Road, Flanagan 2193. Somerset

East: Zuurberg, Annsvilla, Long 952. Uitenhage:

Swartkops River, Zeyher 915 (NBG, S).

Transvaal. — Petersburg: Brak River, Breme-

kamp & Schweickerdt 39; Rapetse, Schlechter

4673. Soutpansberg: Duvenage’s Pan, near Amis-

fort, Schweickerdt & Verdoorn 625.

S.W.A. — Gibeon: Packriem, Range 1361. Groot-

fontein: Otavi, Dinter 5477; Okongawa, Granit-

bank, Dinter 6966. Kaokoveld: Kowares, Hall

343. Outjo: Between Outjo and Otjikondo, Barnard

in SAM 33299. Rehoboth: Buellsport, Strey 2080.

Warmbad: Great Karasberg — Naruda Slid, Keiop

River, Pearson 8456. Welwitschia: near Ugab

River Bridge, Hardy & De Winter 1495. Windhoek:

Neudam Experimental farm, Van Vuuren 1022.

Although the type specimen, Zeyher 191, could

not be located, there were two Zeyher collections

in the S.A. Museum Herbarium (Nos. 912 and 4305)

which were named A. desertorum Spreng. by Zeyher

himself. The description of the type is very short

but it does mention that the leaves are cordate,

which make it undoubtedly this species.

Mentioned under the Speke & Grant Expedition,

Botany, in Trans. Linn. Soc. 29 : 158 (1875) is

Aponogeton leptostachyus var. minor Bak. It is

based on Grant 1863, from Ganga Thembo in the

Sudan. With its bright purple flowers, it may be

a synonym of A. abyssinicus Hochst. Engler, in

his monograph (Bot. Jahrb. 8 : 270, 1887), regards

A. abyssinicus as a variety of A. desertorum, but

they appear to be distinct.

Common name in Rhodesia: Dog with Two Tails.

5. Aponogeton stuhlmannii Engl. inNotizbl.

Bot. Gart. Berlin 1 : 26 (1895); A. Benn. in

F.T.A. 8 : 218 (1901); Krause & Engl, in

Pflanzenr. 4, 13 : 12, t.9 (1906). Type: East

Africa, Usinja, Bugando, Stuhlmann 3541 (B,

holo. !, PRE, photo.).

A. gracilis Schinz ex A. Benn. in F.C. 7 : 43 (1897);

A. gracilis Schinz in Dur. & Schinz, Consp. FI.

Afr. 5 : 492 (1895), nom. nud.; in Bull. Herb.

Boiss, ser. 2, 1 : 736 (1901); Krause & Engl, in

Pflanzenr. 4, 13 : 19, t.9 (1906). Type: Transvaal,

Pietersburg, Houtbos, Rehmann 5761 (K, holo.).

Aponogetonaceae

91

92

Aponogetonaceae

Small tuberiferous herbs 3-12 cm high,

with the upper parts fugaceous. Tubers

developing new plants laterally, globose, up

to 1 cm in diam. brown, densely covered above

with long, fine, white matted roots. Leaves:

the primary (representing the petioles only)

submerged, awl-shaped falcate, about 2 cm

long, 1 mm in diam., present in young seed-

lings; later leaves forming a long filiform

petiole and a floating lamina, linear-oblong,

2-4 cm long and 5-6 mm broad, apex and base

obtuse. Spikes simple, about 9-flowered,

short at first, elongating in fruit, up to 2 cm

long; spathe spotted, membranous; flowers

at first dorsiventrally arranged in 2 rows, after-

wards the rhachis twists in a zig-zag fashion

and the arrangement then appears more

cylindrical; perianth-segments 2, white, ob-

long, 5 mm long, deciduous, apex obtuse or

emarginate. Stamens about 6, filaments nar-

rowly subulate, 1-2 mm long, anthers yellow,

turning black. Carpels c.3, whitish to purplish

or bluish, transparent, with 4-6 basal ovules ;

style about as long as the ovary. Follicles

turbinate, 1-4 seeded; seeds black, shiny,

fusiform, 2 mm long. Fig. 27 : 2.

Recorded from the Transvaal and South West

Africa; also in East Africa and Rhodesia. Usually

found in granite rockpools that are dry most of the

year. A fugaceous plant developing very fast under

favourable conditions, the tuber becoming dormant

again when the pool dries up.

Transvaal. — Bronkhorstspruit: farm Roode-

poortjie 149, Repton 4714a, b. Pietersburg: Hout-

bosberg, Schlechter 4707. Soutpansberg: Dongola,

Greefswald, Pole Evans 4523; Verdoorn 2268;

farm Zoutpan 193, Obermeyer, Schweickerdt &

Verdoorn 317a.

S.W.A. — Otjiwarongo: Waterberg Plateau, Boss

in TRY 35082.

JUNCAGINACEAE

by A. A. Obermeyer

Perennial or annual, glabrous, scapose marsh herbs or aquatics. Roots fibrous. Rhizome

stoloniferous or bulbous, often covered with fibres from old leaf-bases. Leaves basal, dis-

tichous, rush-like, erect or occasionally floating, linear to cylindrical, forming a broad open

basal sheath; axillary scales present. Inflorescence a simple raceme or spike, ebracteate;

flowers anemophilous, protogynous, bisexual or unisexual and dioecious or polygamous,

actinomorphic or somewhat irregular, perianth-segments 6-2, the inner whorl raised above

the outer. Stamens 6-4, anthers sessile or subsessile, epipetalous. Carpels 6-3, free or connate,

uniovulate, ovule erect or pendulous. Fruit with 6 carpels or usually fewer by abortion, dry,

cylindrical, ovoid, obovoid or globose, free or, if connate, separating upwards from a central

axis. Seed exendospermous.

A small cosmopolitan family, well represented in Australia, frequenting saline or fresh-water habitats.

The chief genus, Triglochin, is the only one rich in species and is widely distributed, being the only genus

occurring in Southern Africa. Two monotypic genera, Maundia and Tetroncium are confined to Australia,

the Antarctic and South America. Hutchinson places Scheuchzeria, also a monotypic genus from the North

Temperate Zone, in a family by itself, the Scheuchzeriaceae. Other taxonomists, e.g. Buchenau, Marloth,

Adamson, etc., unite the two families but, in this case, the name Juncaginaceae, being older, must get priority.

Lilaea, another monotypic genus from America, included in this family by Bentham and Hooker, Rendle

and others, is best placed in a separate family, the Lilaeaceae, according to more recent authors, who take

into consideration the conclusions of Uhl’s research on Helobiae.

66 TRIGLOCHIN

Triglochin L., Sp. PI. 338 (1753); Gen. PI. ed. 5 : 157 (1754); Micheli in DC., Monogr. Phan.

3 : 96 (1881); Buchen. in Bot. Jahrb. 2 : 490 (1882); Benth. & Hook.f., Gen. PI. 3 : 1012 (1883);

JUNCAGINACEAE

93

A. Benn. in F.C. 7 : 41 (1897); Rendle in Cat. Afr. PI. Welw. 2 : 93 (1899); A. Benn. in F.T.A.

8:215 (1901); Buchen. in Pflanzenr. 4, 14 : 7 (1903); Phill., Gen. ed. 2 : 56 (1951); Adamson

in FI. Cape Penins. 38 (1950); Rendle in Class. Flow. PI. ed. 2, 1 : 208 (1953). Type species:

T. palustris L.

Herbaceous scapose marsh plants, perennial (annual in some Australian spp.). Roots

fibrous with a spongy outer covering that soon disintegrates and a wiry thin stele; with thin

side roots; root-hairs long. Rhizome stoloniferous or bulbous, covered with fibres from old

leaf-bases. Leaves basal, distichous, rush-like, forming a broad open, ligulate sheath below,

cylindrical to setaceous, glaucous, somewhat succulent; axillary scales minute, linguiform.

Inflorescence a simple raceme overtopping the leaves; scape terete, firm; flowers ebracteate

on decurrent, erecto-patent pedicels, actinomorphous, perianth segments 6, green, the 3 inner

raised above the outer whorl, conchiform with the 6 sessile anthers inserted inside the hollow

segments (occasionally some stamens aborted in species outside South Africa), locules extrorse,

dorsifixed, dithecous, dehiscing longitudinally, pollen globose; the segment and the stamen

semi-persistent, falling together; carpels 6 or the outer whorl suppressed, connate, styles

plumose, sessile or subsessile, the solitary ovule erect. Follicles cylindrical, globose, ovoid, or

obovoid sometimes spurred below, separating upwards from a central columella. Seed erect,

embryo straight.

A genus of about 14 species, cosmopolitan (about half of these Australian). Two species found in Southern

Africa, in salt marshes, on stream banks and mountain slopes.

“The generic name Triglocliin was treated as neuter by Linnaeus and by many authors following him. But

the word is a compound of the Greek Tri- (three) and glochin (projecting point), and as glochin is feminine,

the name Triglochin is to be treated as feminine”. J. E. Dandy, in a personal communication.

Follicle elongate, 5-10 mm long with 3 fertile carpels and no sterile carpels 1 . T. bulbosa

Follicle rounded, 2 mm long with 3 fertile carpels alternating with 3 sterile ones 2. T. striata

1. Triglochin bulbosa L., Mant. 2 : 226

(1771); Jacq., Ic. Rar. 2, t.454; Coll. Suppl.

102 (1786-1793); Thunb., Prodr. 67 (1794);

FI. Cap. ed. Schult. 2 : 347 (1823); Ker

Gawler in Bot. Mag. t.1445 (1812); Micheli

in DC., Monogr. Phan. 3 : 99 (1881); Buchen.

& Hieron. in Pflanzenfam. 2 : 224 (1889); A.

Benn. in F.C. 7 : 42 (1897); F.T.A. 8 : 215

(1901); Buchen. in Pflanzenr. 4, 14:11 (1903);

Marloth, FI. S. Afr. 4 : 14, t.2, fig. 5 (1915);

Adamson in FI. Cape Penins. 39(1950). Type:

Cape (LINN 466-3; PRE, photo.).

T. elongata Buchen. in Pflanzenr. 4, 14 : 10,

fig. 3 (1903). Syntypes: Cape, Malmesbury, near

Hopefield, Bachmann 1692; 1693 (Bf); Pondoland,

Bachmann 309 (Bf). T. laxiflora sensu A. Benn. in

F.C. 7 : 42 (1897), as to Rehmann 8581, Wood 925;

sensu Buchen. in Pflanzenr. 4, 14 : 11 (1903), as

to Schlechter 10424; sensu Adamson in FI. Cape

Penins. 38 (1950). T. tenuifolia (sphalm. tenuifolium )

Adamson in Journ. S. Afr. Bot. 5 : 30 (1939); FI.

Cape Penins. 39 (1950). Type: Cape, Table

Mountain, Adamson 968 (BOL, holo.). T. compacta

(sphalm. “ compactum ”) Adamson in Journ. S. Afr.

Bot. 9 : 152 (1943); FI. Cape Penins. 39 (1950).

Type: Cape, near Houtbay, Compton 14436 (NBG,

holo.). T. milnei Horn af Rantzien in Svensk. Bot.

Tidskr. 55 : 85 (1961). Type: Zambia, Mwinilunga,

Milne-Redhead 3012 (K, holo.). T. bulbosa L. subsp.

tenuifolia (Adamson) Horn af Rantzien in Svensk

Bot. Tidskr. 55 : 106 (1961).

Perennials, variable in size ; when crushed

emitting a pungent smell (like a Jerusalem

Artichoke). Roots fibrous and with several

larger contractile roots. Rhizome stoloniferous

or cormous (when the stolons are arrested in

hard soils), often forming cormlets, the fibrous

covering hard and sclerotic in dry surround-

ings or soft or nearly absent in moist con-

ditions. Leaves variable in size, 5-35 cm long

and 0-5-2 mm broad, filiform to semi-terete,

with a shallow ventral groove, gradually

tapering to an acicular apex, expanded below

to form an open, semi-amplexicaul sheath,

glaucous, succulent; the primary leaves, if

present, bract-like, scaly, brown. Inflorescence

an erect central, few to many flowered raceme ;

94

JUNCAGINACEAE

scape terete, firm; pedicels patent to erecto-

patent, decurrent, lengthening in fruit, 2-10

mm long; flowers green with the apiculate

tips of the segments often purplish, segments

conchiform, 2-3 mm long; stigmas 3, papil-

lose, purple, sessile, 3 outer carpels vestigial,

3 inner soon lengthening at anthesis and after

fertilization. Follicles 5-10 mm long, narrow-

ly ovoid-acuminate to cylindrical, tipped by

the 3 persistent, hook-like style-bases, with a

very short spur below. Fig. 28 : 1.

Widely distributed in Southern Africa, extending

to tropical Africa and the Mediterranean; common

along the coast in salt marshes and bordering

lagoons; the typical (cormous) form is found on

mountain slopes in non-saline habitats.

Cape. — -Bathurst: Alexandria, Dyer 3365. Cal-

vinia: Lokenburg, Acocks 17470. Ceres: Ceres

Road (Wolseley), Schlechter 8979. East London:

Kwenqura River Mouth, Galpin 5815. George:

Kaaimans River, Wilman. Kentani: Mazeppa Bay,

Story 4470. Malmesbury: Mamre Hills, Barker

1596. Oudtshoorn: De Rust, Acocks 20454. Penin-

sula: Karbonkelberg, Leighton 446; sandy shore

towards Camps Bay, Wolley Dod 3459; Diep River,

Wolley Dod 2573. Robertson: Bonnievale, Marloth

11592. Stellenbosch: Koelenhof, Strey 670.

Natal.— Durban : Congella, Wood 11985.

Mtunzini: Richards Bay, MacNae. Umlazi:

Isipingo, Green 37.

The typical form, judging from the (nearly

leafless) type (LINN 466.3), must have come from

a montane habitat. Adamson’s species T. tenuifolia

resembles it closely and must be regarded as a

synonym. Many similar slender plants hailing from

the south-western Cape mountains have the fruiting

pedicels long and patent, but this is not a constant

character although used by many systematists. In

the type plants the pedicels, although immature,

appear to be erecto-patent and fairly short.

Ecklon in Linnaea, p. 1 3 1 (1832), already observes

that the species may grow in saline sand or on

mountain slopes. That the plants can adapt them-

selves to saline or non-saline conditions was proved

by cultivation at the Botanical Research Institute.

In mud the stolons elongate but in firm soils

they are arrested and form corms. These become

covered with the persistent sclerotic fibres of the

leaf bases. The species T. elongata, T. compacta

and T. milnei, based on the differences in the under-

ground parts, must be considered synonyms of

T. bulbosa L. (If Horn af Rantzien had been aware

of Adamson’s species T. compacta , I am sure he

would have referred the material from Zambia and

Natal to this species.) The plants flower freely and

intermittently during summer and produce much

fertile seed. Those growing in moist habitats

become large and coarse, with short erect pedicels,

whereas the montane form remains small and

slender. T. palustris L., from Europe, is said to

make temporary bulbs (winterbuds) in autumn.

The formation of storage organs in T. bulbosa may

serve a similar purpose. Both the stoloniferous

and cormous forms, the latter also with cormlets,

are present in Wolley Dod’s gathering (No. 2573)

from the Cape Peninsula.

2. Triglochin striata Ruiz & Pavon , FI.

Peruv. et Chil. 3 : 72 (1802); Micheli in DC.,

Monogr. Phan. 3 : 101 (1881); A. Benn. in

F.C. 7 : 42 (1897); Rendle in Cat. Afr. PI.

Welw. 2 : 93 (1899); A. Benn. in F.T.A.

8 : 216 (1901); Buchen. in Pflanzenr. 4, 14 : 10

(1903); Adamson in FI. Cape Penins. 38

(1950). Type: South America.

T. maritima sensu Thunb., Prodr. FI. Cap.

1 : 67 (1794); FI. Cap. ed. Schult. 2 : 340 (1823),

non L. — var. /?. micrantha E. Mey., Linnaea

7 : 131 (1832). T. natalensis Gand. in Bull. Soc. Bot.

France 66 : 294 (1919). Type: Durban, Schlechter

2857 (K).

Perennials, variable in size, 5-50 cm high,

forming long, woody stolons. Roots firm and

coarse interspersed with thin young matted

feeding roots. Rhizome woody, sparsely

covered with fibres. Leaves arranged in a

compact fan-shaped manner with the sheaths

broad, distinctly ligulate, lamina cylindrical,

5-30 cm long, succulent, green, apex obtuse.

Inflorescence ultimately overtopping the

leaves, the flowers arranged in dense _.^>al

whorls, pedicels arcuate, decurrent, very short

at first, up to 3 mm long in fruit. Flowers

small, segments 1 mm long; carpels 6, the 3

inner fertile alternating with the 3 sterile

outer. Follicles globose, 2 mm in diam.,

3-costate, compressed, shortly stipitate, apicu-

late. Fig. 28 : 2.

Recorded from the temperate zones of the

Americas, Australia, New Zealand and Southern

Africa, extending to Angola in the west and Mozam-

bique in the east; occurs in saline marshes and

borders of lagoons near the sea, often in mangrove

swamps.

Cape. — Bathurst; Kowie River Lagoon, Mauve &

Wells 8. Caledon: Hermanus, Rogers 26576.

Clanwilliam: Graafwater, Acocks 19778. East

London: Galpin 7353. Humansdorp: Kabeljous

River mouth. Mauve & Wells 56. Knysna: Ruigte-

vlei, Martin 4541. Peninsula: Noordhoek salt pan,

Tdlken 442; Ronde Vlei, Andreae 238. Port

Elizabeth: Chatty River, Mauve & Wells 45. Port

St. Johns: Wager. Riversdale: Gouritz River

Mouth, Tdlken 353. Willowvale: Bashee River

Mouth, Macnae.

Natal. — Durban: De Winter 7718. Hlabisa:

St. Lucia Estuary, Ward 3392. Ingwavuma: Kosi

Estuary, Tinley & Ward 12. Mtunzini: Richards

Bay, Macnae. Ubombo: Sordwana Bay, Tinley 452.

JUNCAGINACEAE

95

Fig. 28. — 1, Triglochin bulbosa, x 1 ; la, base of plant with stolons, x 1 ; lb, flower, x 6; lc, follicles, x 5;

Id, follicle, axial view, x 5. 2, T. striata, X 1; 2a, follicles, X 10.

96

ALISMATACEAE

by A. A. Obermeyer

Alismataceae

Perennial, usually deciduous, occasionally annual, aquatic or marsh herbs, lactiferous.

Roots thin, abundant, branched. Rhizome woody and irregular or tuberous, suckering or

forming runners bearing winterbuds. Leaves basal, subdistichous, with the usually long

petioles expanded below to form an open sheath; blade erect, entire, lanceolate, ovate

hastate or sagittate, rarely floating and/or submerged and then phyllodic; with distinct

parallel veins converging to the apex and lobes, connected by cross-veins; with an apical

hydathode; axillary scales present. Inflorescence usually a tall pedunculate compound, usually

trimerous, raceme or panicle, reduced in some species to one flower. Flowers bisexual and/or

unisexual (rarely dioecious) regular, bractiferous; sepals 3, green, persistent; petals 3, delicate,

fugaceous. Stamens 3-oc, free; filaments filiform or expanded below; anthers bilocular,

basifixed, pollen sphaeroid, pollination entomophilous, occasionally cleistogamous; stami-

nodes present in some species. Carpels 2-oc, free (or united at the base in Damasonium, a

genus outside South Africa), if numerous situated on a flat or convex receptacle, arranged

spirally or in a ring, unilocular, the style terminal or ventral, stigma apical, uni-ovulate

(pluri-ovulate in Damasonium ); in unisexual male flowers sterile carpels may be present.

Fruit a follicle, drupe or achene, buoyant; seeds with a horseshoe-shaped embryo,

exendospermous.

Predominantly a family of the northern hemisphere, in temperate and tropical regions, with 13 genera

and about 70 species. In Southern Africa 3 genera are found, each represented by one widespread species.

Leaves linear-lanceolate to ovate, base cuneate to cordate:

Plants dioecious; petals minute or absent; achenes irregularly crowded on the

receptacle 3. Burnatia

Plants monoecious; petals larger than the sepals; achenes neatly arranged in a circle

on the receptacle 1 . Alisma

Leaves hastate 2. Limnophyton

70 1. ALISMA

Alisma L., Sp. PI. 342 (1753); Gen. PI. ed. 5 : 160 (1754); Micheli in DC., Monogr. Phan.

3 : 31 (1881); Benth. & Hook.f., Gen. PI. 3 : 1004 (1883); Wright in F.T.A. 8 : 207 (1901);

Buchen. in Pflanzenr. 4, 15 : 12 (1903); Carter in F.T.E.A. Alismataceae: 5 (1960). Type

species: A. plantago-aquatica L.

Aquatic scapose herbs, perennial from a woody, irregular rhizome, suckering. Roots

fine, very abundant. Leaves erect, blade variable, linear-lanceolate to elliptic, base rounded

or cordate. Inflorescence a verticillate, pyramidal, compound panicle or often reduced.

Flowers pedicellate, bisexual; stamens 6, in pairs opposite the petals, filaments filiform;

carpels 10-20, free, situated in a circle on the torus, compressed laterally, style ventral, short,

filiform; ovule campylotropous. Achenes many, compressed, sulcate.

Cosmopolitan; considered monospecific by some, subdivided into several species and subspecies by

others. One species recognized in Southern Africa.

1. Alisma plantago-aquatica L., Sp. PI. Carter in F.T.E.A. Alismataceae: 5 (1960);

342 (1753); Wright in F.T.A. 8 : 207 (1902); Wild in Kirkia 2 : 36, t,13a (1961). Type:

Buchen. in Pflanzenr. 4, 15 : 13 (1903); Europe (LINN, 473 • 1). Chromosome num-

Adamson in FI. Cape Penins. 39 (1950); ber 2n = 14.

Alismataceae

97

Perennial, stoloniferous, scapose, up to

1 m high. Roots thin, densely covering the

rhizome. Leaves with erect petioles up to

40 cm long; lamina ovate, rounded to cordate

at the base, usually about 10 cm long and

6 cm broad (larger in tropical specimens),

nerves usually 5, laterals closely parallel to

one another, ascending. Inflorescence a much

branched pyramidal panicle (occasionally

poorly developed); peduncle about 50 cm

high (taller in tropical specimens), terete;

panicle with about 6 whorls, the lower com-

pound, ultimately bearing 1-6 pedicelled

flowers in terminal umbels; bracts scarious,

acuminate, lowest 25 mm long, younger be-

coming progressively smaller, deciduous;

pedicels up to 2 cm long; sepals ovate, green,

2 mm long; petals white or pink; carpels

about 20. Achenes ovate, 2 mm long, com-

pressed laterally, deeply furrowed dorsally.

Fig. 29 : 1.

Cosmopolitan. Apparently a recent adventive

in South Africa, recorded from one area near

Cape Town and from the Transvaal highveld, the

Vaal River and its tributaries ; common where found.

Cape. — Peninsula: near Cape Town, Moss 4185.

O.F.S. — Vredefort: Parys, Obermeyer in TRV

31673.

Transvaal. — Heidelberg: Alberton, Dimovic in

J28023. Johannesburg: Moss 8404. Potchefstroom :

Nooitgedacht, Vaal River, Louw 1640.

The seeds are buoyant because of subepidermal

air-tissue and may float for many months. Eaten

by waterbirds, the seeds pass unharmed through

the alimentary canal.

Common name: Water-plantain.

72 2. LIMNOPHYTON

Limnophyton Miq., FI. Ind. Bat. 3 : 242 (1855); Benth. & Hook.f., Gen. PI. 3 : 1005 (1833);

Wright in F.T.A. 8 : 209 (1902); Buchen. in Pflanzenfam. 2, 1 : 231 (1889); Pflanzenr.

4, 15 : 21 (1903); Phill., Gen. ed. 2 : 56 (1951); den Hartog in FI. Males. 1, 5 : 324(1957);

Carter in F.T.E.A. Alismataceae: 7 (1960). Type species: L. obtusifolium (L.) Miq.

Aquatics or marsh herbs. Leaves hastate, glabrous or pubescent. Panicle pyramidal,

whorled; flowers bisexual and male in lower whorls, male only in upper; sepals 3, boat-

shaped, reflexed after anthesis; petals 3, white, orbicular, clawed; stamens 6, filaments

bulbous below; carpels numerous, crowded on a small globose receptacle, style ventral,

ovule solitary, basal. Achenes asymmetrical, globose, shortly stipitate, with 2 lateral air-

chambers between the exocarp and endocarp.

Three species of which one occurs in Southern Africa and is widespread in tropical Africa and south-east

Asia ; the other 2 are endemic in tropical Africa. Found in or beside fresh water pools or slow-flowing rivers.

Limnophyton obtusifolium ( L.)Miq ., FI.

Ind. Bat. 3 : 242 (1855); Wright in F.T.A.

8 : 209 (1902); Buchen. in Pflanzenr. 4,

15 : 22, fig. 10 (1903); den Hartog in FI.

Males. 1, 5 : 324 fig. 4 (1957); Carter in

F.T.E.A., Alismataceae: 9 (1960). Type:

Asia, Plukenet, t.220, fig. 7 in Herb. Sloane,

Vol. 97, fol.181 (BM, lecto.).

Sagittaria obtusifolia L., Sp. PI. 993 (1753).

Apparently annuals, perennating occa-

sionally, up to 1 m high. Roots fibrous,

spongy, with transverse septa. Rhizome

small. Leaves on long petioles which are

triangular in cross-section, spongy, the trans-

verse septa distinct in juveniles; lamina erect,

hastate, with the angle between the basal lobes

obtuse (more than 90°), the lobes small at

first, about as long as the upper part of the

blade in mature leaves, apex obtuse, nerves

13-15, the 3-4 lower curving downwards,

glabrous or glabrescent, pellucid dotted.

Inflorescence a pyramidal panicle up to 1 m

high when fully developed, often much re-

duced in young or small plants; scape stout,

woody ; flowers in 3-7 dense whorls the lower

compound; bracts scarious; lanceolate-

acuminate, reflexed with age; fruiting pedicels

up to 15 mm long, swollen and woody, sterile

pedicels wiry. Achenes about 30, forming a

globose head 12 mm in diam., each fruit hard,

brown, obpyramidal, 5 mm long, shortly

stipitate, ridged. Fig. 29 : 2.

Widespread in tropical Africa, Madagascar and

south-east Asia. Recorded from the Transvaal

and Natal at low altitudes; common where found.

Natal. — Ubombo: Mkuzi Game Reserve, Ward

3070; Tinley 618.

98

Alismataceae

r v.d n

Fig. 29.— 1, Alisma plantago-aquatica, x £; la, fruit, x 5; lb, seed, x 10. 2, Limnophyton obtusifolium,

X i; 2a, seed. 3, Burnatia enneandra, X 3a, fruit, x 2; 3b, seed, X 12.

Alismataceae

99

Transvaal. — Pilgrim’s Rest: Kruger National

Park, near Skukuza, Code I 6071; Tshokwane,

Leeupan, Van der Schijjf 2765 ; 4194. Soutpansberg:

Dongola Reserve, Schroda Lily Pool, Pole Evans

4587.

It is possible that L. angolense Buchen. may

occur in the northern region of South West Africa.

It can be distinguished from L. obtusifolium by

being more hirsute, by its long-attenuate bracts

its longer and more slender pedicels and its achenes

which are longitudinally ridged when dry; the angle

between the lobes of the leaf at the base is acute.

The inflorescence too appears more cylindrical, as

the lower side branches remain shorter. The syn-

types are Baum 364 from Kubango, and Mechow

from Malange, Angola.

77 3. BURNATIA

Burnatia Micheli in DC., Monogr. Phan. 3 : 81 (1881); Benth. & Hook.f., Gen. PI. 3 : 1007

(1883); Wright in F.T.A. 8 ; 212 (1901); Buchen. in Pflanzenr. 4, 15 : 59 (1903); Carter in

F.T.E.A. Alismataceae: 13 (1960). Type species: B. enneandra Micheli.

Rautanenia Buchen. in Bull. Herb. Boiss. 5 : 855 (1897); Pflanzenfam. Nachtr. 2 : 3 (1900); Pflanzenr.

4, 15 : 59 (1903); Phill., Gen. ed. 2 : 56 (1951).

Perennial, aquatic, scapose herbs. Rhizome bulbous covered with short thin roots.

Leaves erect, on long petioles, blades linear to lanceolate, the base decurrent to rounded.

Inflorescence overtopping the leaves, a trimerous compound umbel, sometimes reduced and

few-flowered ; with 3 basal acuminate bracts which at first envelope the umbel in bud, scarious,

semi-persistent; upper bracts reduced or absent. Flowers 3 in a whorl, evenly spaced, pedi-

celled; male flowers with 3 concave sepals, petals 3, smaller than the sepals, stamens 9;

sterile carpels about 12; female flowers with 3 concave sepals, petals 0 or minute, staminodes

0-1-2; carpels many, closely arranged on a small receptacle, compressed, stigma ventral,

discoid, papillose, subsessile, ovule basal, solitary. Achene obovate-orbicular laterally com-

pressed, with 2 narrow circular wings.

Monospecific, confined to tropical and subtropical

or marshes, the parts above ground dying down after

Burnatia enneandra Micheli in DC.,

Monogr. Phan. 3 : 81 (1881); Buchen. in

Pflanzenfam. 2 : 229, 232 (1889); Pflanzenr.

4,15: 60 (1903). Type: Sudan, Kordofan,

Kotschy 192 (BM, K).

Echinodorus ? schinzii Buchen. in Bull. Herb. Boiss.

4 : 413 (1896). Type: South West Africa, Ovambo-

land, Omulonga, Rautanen 51 (Z, holo.)-

Rautanenia schinzii (Buchen.) Buchen. in Bull.

Herb. Boiss. 5 : 854 (1897); Wright in F.T.A.

8 : 212 (1901); Buchen. in Pflanzenr. 4, 15 : 59

(1903).

Scapose herbs up to 1 m high. Leaves

wholly or partially submerged with a flattened

petiole up to 60 cm long, gradually widening

to form a linear to lanceolate lamina above,

up to 18 cm long and 3 cm broad but very

variable. Male inflorescence forming a deli-

cate, finely branched pseudo-umbel with the

small flowers on filiform pedicels up to 1 cm

long; sepals orbicular, concave, 4 mm long,

delicate pale green to purplish with trans-

parent margins; petals 1-2-5 mm long,

narrow; stamens included, white; sterile

Africa; forming clumps in or near temporary pools

flowering.

carpels light green. Female inflorescence

smaller than the male with fewer branches

and the flowers in trimerous subsessile clus-

ters; sepals delicate, orbiculate, concave,

1-5 mm long; petals absent (present as

minute scales in robust specimens, vide

Carter); carpels about 12 on a short cylindri-

cal torus, compressed and with 2 lateral ear-

shaped or horseshoe-shaped rims or folds.

Achenes black, 1-5 mm long. Fig. 29 : 3.

Widespread in tropical Africa. Recorded from

northern South West Africa and the Caprivi Strip

where it is locally common.

S.W.A. — Okavango Native Territory: Okavango

River Flats between Runtu and Mupini, Marais

1114. Grootfontein : near Andara, Merxmii/ler

2088. Caprivi: Katima Mulilo area, Killick &

Leis tner 3113.

Sweet smelling, like Eau de Cologne.

Carter in F.T.E.A. Alismataceae: 15 (1960) also

enumerates as synonyms Burnatia alismatoides

Peter and its variety elliptica Peter, B. enneandra

Mich. var. linearis Peter and B. oblonga Peter, all

from Tanganyika. The types were preserved at

Berlin, but have been destroyed.

100

HYDRO CHARIT ACE AE

by A. A. Obermeyer

Hydrocharitaceae

Fresh-water or marine, annual or perennial herbs, partially or completely submerged.

Roots terrestrial or floating. Leaves radical, in rosettes or cauline, alternate, opposite or in

verticils, submerged or with floating blades. Flowers submerged in marine genera, in others

exserted during anthesis, monoecious or dioecious, unisexual or rarely bisexual; usually

exserted from a pedunculate, or rarely sessile, tubular, bind spathe, or rarely subtended by 2

free, opposite bracts; male flowers usually numerous and, in some genera, become detached

and free floating; female flowers mostly solitary inside the spathe. Male flowers with perianth

regular, rarely irregular ( Vallisneria ), 1-2-seriate with the 3-2 outer segments often sepaloid,

valvate, and the 3-2 inner petaloid, imbricate; stamens 3-many (rarely 1), in one or more

trimerous whorls, the inner occasionally staminodial; anthers with parallel locules, opening

longitudinally; rudimentary ovary sometimes present; pollen sphaeroid or, in marine genera,

confervoid or united in strings; pollination in marine genera hydrophyllous, in others ento-

mophilous. Female flowers with an inferior, 1-celled ovary, with 3-6 or rarely more parietal

placentas which at times protrude towards the centre; number of styles equalling the number

of placentas; stigmas simple or bifid. Fruit rounded or tubular, fleshy or leathery, usually

rostrate with persistent style-bases, the seeds being dispersed by the dissolution of the walls

which become mucilaginous; rarely fruits dry and dehiscent. Seeds exendospermous, with a

large embryo.

A family consisting of 14 divergent genera, several of which are monotypic, and about 80 species. Three

are marine with a mainly tropical and subtropical distribution, while the others are found in fresh water,

widely distributed both in tropical and temperate regions. In Southern Africa one marine and two fresh-

water genera, comprising 10 species, have been recorded.

Egeria densa Planch., a South American species, sometimes wrongly identified as Elodea or Anacharis,

has become an obnoxious adventive in New Zealand and has been introduced into Natal, where it should

be carefully watched. So far no records of it becoming naturalised have come to light, but it is cultivated

in many ponds in parks, universities, gardens and in aquaria. A large pond on the Tongaat Sugar Estates

has to be cleared regularly to allow other aquatics, such as Nymphaeas, to grow and flower. It was probably

imported in Medley Wood’s time when a submerged aquatic was needed as an “oxygen plant” for the

fishes introduced to combat malaria-carrying mosquito larvae. Only male plants were imported (females

being rare) but they propagate at an alarming rate. The species resembles Lagarosiphon major but is coarser

and has the leaves in whorls of 2-5. The yellow or cream-coloured male flowers remain attached to the plant

and are entomophilous. The perianth forms a long filiform tube with the 3 inner segments forming a cup

about 1 cm in diameter, which is exserted above the surface. There are 9 stamens.

The genus Hydrilla Rich., wrongly included by Phillips, Gen. ed. 2 : 57 (1951), is absent from Southern

Africa.

Marine plants; leaves linear to lanceolate-ovate, in pairs from a creeping rhizome; flowers

apetalous, submerged 1. Halophila

Fresh-water plants; leaves rosulate or cauline; flowers with petals, exserted above water

during anthesis:

Forming long leafy submerged stems from a perennial rhizome; spathes sessile,

axillary 2. Lagarosiphon

Rosulate, acauline annuals; spathes pedunculate 3. Ottelia

85 1. HALOPHILA

Halophila Thouars, Gen. Nov. Madag. 2 (1808); Aschers. & Guerke in Pflanzenfam. 2, 1 : 247

(1889); Wright in F.T.A. 7 : 10 (1898); Phill., Gen. ed. 2 : 57 (1951); den Hartog in FI. Males.

1,5:407 (1958). Type species: FI. stipulacea (Forsk.) Aschers.

Hydrocharitaceae

101

Marine perennials with a much-branched creeping rhizome. Roots simple, 1 or rarely

2 from a node, densely covered by long root-hairs. Rhizomes bearing at the nodes an

abbreviated shoot and a pair of leaves, each surrounded by a basal scale. Leaves opposite,

sessile or petiolate, linear, lanceolate, oblong or ovate, entire or serrulate, glabrous or

pubescent; midrib distinct, with a prominent lateral nerve next to the margin, which is

nnected to the midrib by many oblique secondary nerves. Flowers monoecious or dioecious,

unisexual, usually solitary in the axil of a secondary shoot, rarely 1-2 male and a female

flower together in a spathe; spathe of 2 free, membranous bracts. Male flowers shortly

pedicelled, with 3 perianth segments; stamens 3, anthers sessile, 2-4 celled, extrorse; pollen

adhering to form threads, pollination hydrophyllous. Female flower sessile; ovary ovoid,

1-celled, beaked, bearing 3 reduced perianth segments near the apex; styles 2-5, linear.

Fruit ovoid, rostrate, thin-walled. Seeds few to many, small, globose.

Found in Africa, Asia, Australia and America; there are 9 species of which some are widely distributed,

others endemic. One widespread species occurs in Southern Africa.

Halophila ovalis ( R.Br .) Hook.f., FI.

Tasm. 2 : 45 (1858); Aschers. in Linnaea

35 : 173 (1868); Balf.f. in Trans. Bot. Soc.

Edinb. 13 : 290, t.2, fig. 11-13, 17, etc. (1879);

Aschers. & Guerke in Pflanzenfam. 2, 1 : 249,

fig. 182 (1889); Setchell in Bull. Torr. Bot.

Club 47 : 570 (1920) ; Kausik & Rao in Journ.

Mysore Univ. 3 : 41 (1942); Perrier de la

Bathie, FI. Madag. fam. 26 : 4, fig. 1, 3-5

(1946); Macnae in Nat. Hist. Inhaca Island,

Mozambique 28, fig. 8b, c (1958); den Hartog

in FI. Males. 1,5: 408 (1958); Subramanyam,

Aquatic Angiosperms 62, t.44 (1962). Type:

Australia.

Caulinial ovalis R.Br., Prodr. FI. Nov. Holl.

339 (1810).

Halophila ovata sensu Wright in F.T.A. 7 : 10

(1898). H. linearis den Hartog in Acta Bot. Neerl.

6 : 46 (1956). Type: Mozambique, Inhaca Island,

Cohen in Herb. Moss 20652 (BM, holo.).

Fig. 30. — 1, Halophila ovalis, x 1; a, male flower, x 1; b, female flower, x 1; c, linear leaf, X 1.

102

Hydrocharitaceae

Dioecious. Rhizomes long, branching;

internodes about 2-4 cm long, light yellow,

thin, brittle. Leaves in pairs, subtended by

bracts; petiole 1-3 cm long; lamina variable

in shape and size, linear to lanceolate or

ovate (“racquet-shaped”), 10-15 mm long

and 2-8 mm wide, green often with brown

transverse lines, apex acute, or rounded, base

attenuate or rounded. Male flowers shortly

pedicellate, with 3 concave perianth segments,

4 mm long and 2 mm broad ; anthers oblong,

2-4 celled. Female flowers enclosed in 2 con-

cave, ovate bracts, consisting of an ovoid,

beaked ovary 4 mm long, terminating in 3

styles 10-20 mm long, soon deciduous. Fruit

globular, 4 mm in diameter with the beak

persistent. Seeds c. 20, rounded, 1 mm long,

tuberculate, reticulate. Fig. 30.

Occurs along the eastern coast as far south as

Knysna Lagoon, submerged in from 1-30 ft. of

water, nearly or just exposed at very low tides.

A widespread, common species found along the

coast of eastern Africa and the Red Sea to Indonesia,

Japan, Hawaii, Australia and Tasmania.

Natal. — Durban Bay, western mud-bank near

Maydon Wharf, Ward 451

Cape.— Bathurst: Kowie West, Britten 2663;

5204; Kenton-on-Sea, Bushman’s River Mouth,

Mauve & Wells 17 (with d and $ flowers, January,

1964). Knysna: Lagoon, near Brenton, Duthie

1215; Lagoon west of Leisure Isle, Mauve & Wells

62 (with and $ flowers, January, 1964).

In southern Africa (including Mozambique) the

plants seen were just covered at low spring tides.

From other parts of the world they are reported

from a depth of up to 10 metres. Flowers are rare.

The narrow-leaved form described by den Hartog

as H. linearis (from Mozambique) may be a variety

of H. ovalis. Like most aquatics, multiplication is

mostly vegetative and, as a result, homogeneous

colonies are developed. As the narrow-leaved form,

however, grows together with or near the wide-

leaved form (at least in southern Africa), it seems

unlikely that it could be a separate species. Inter-

mediates do occur and no other differences could be

detected.

88 2. LAGAROSIPHON

Lagarosiphon Harv. in Joum. Bot. Lond. 4 : 230 (1841); Benth. & Hook.f., Gen. PI. 3 ; 450

(1883); Wright in F.T.A. 7 : 2 (1897); F.C. 5, 3 : 1 (1912); Phill., Gen. ed. 2 : 58 (1951);

Oberm. in Bothalia 8 : 139 (1964). Type species: L. muscoides Harv.

Submerged (except for the flowers during anthesis), dioecious, rooted herbs from a

perennial rhizome. Stems long, terete, leafy; branches arising from inside a spathe next to a

flower, the leafy apices compact, the internodes elongating later. Leaves widely spaced below,

dense above, alternate, subopposite or whorled, linear to linear-lanceolate, margin dentate

with unicellular teeth, tissues of the lamina differing in each species (cf. fig. 32), with 2 minute

axillary scales (intra-vaginal squamulae). Male inflorescence axillary; spathes fused, obovate,

ovate or lanceolate, compressed or cup-shaped, dentate; the axis (rudimentary scape) pro-

ducing numerous pedicellate flowers (up to 50 or more) consecutively; the buds in South

African species become detached and rise to the surface (because of an enclosed air bubble)

where they expand and recurve suddenly to form a bell-shaped, floating flower; perianth

with 2 whorls of 3 segments each, the outer slightly narrower; stamens 3, the filaments

ultimately parallel to the water surface with the anthers at right angles to them; each pollen-

sac containing 4 tetrads, grains large; staminodes 3, longer than the stamens, papillate,

usually coloured above, joined at the top (acting as a sail). Female inflorescence axillary;

spathe-valves fused, narrowly oblong, ovoid or cylindrical, entire or toothed, containing 1-3

flowers; perianth-tube exserted laterally near the apex of the spathe, lengthening (meanwhile

giving off gas bubbles to buoy it up) until it reaches the surface of the water (ultimate length

of flower varies with the species); limb 6-lobed; staminodes 3, minute, filiform; ovary

1 -locular with 3 parietal placentas; styles 3, adnate to the perianth-tube, each divided into

2 long, papillate, often brightly coloured stigmatic arms; ovules 6-30, orthotropous, funicle

short or long, straight or bent. Capsule protruding from the torn spathe (the larger inner

spathe-valve bursting along the midrib), rostrate (the persistent perianth-base forming a beak),

ultimately becoming mucilaginous and disintegrating. Seeds cylindrical, shortly stipitate,

apex acute, size varying with each species, closely ribbed or honey-combed, buoyant at first,

sinking later.

Hydrocharitaceae

103

About 16 species occur in Africa; two are endemic to Madagascar; five are found in South Africa, in

fresh water. L. major is reported as an adventive in Europe and New Zealand.

Leaves thin, transparent, 0-5-2 mm broad, cells usually longer than broad:

Leaves with a hyaline margin consisting of at least 3 layers of cells without chlorophyll; the marginal

teeth not on excrescences 1. L. muscoicles

Leaves green to the margin; teeth on small excrescences:

Leaves in verticils of 6-8; central band narrow, not much different from the rest of the lamina

2. L. verticillifolius

Leaves alternate or rarely in verticils; central band usually wide, at least near the base with the

cells much larger and with transverse veins 3. L. crispus

Leaves fairly solid, opaque, 2-3 mm broad, cells small, somewhat papillose-rhomboid:

Teeth of leaves minute, blunt, ascending 4. L. major

Teeth of leaves on triangular excrescences, sharp, patent 5. L. ilicifolius

1 . Lagarosiphon nuiscoides Harv. in Journ.

Bot. Lond. 4 ; 230, t.22 (1842); Wright in

F.T.A. 7 : 3 (1897); F.C. 5, 3 ; 1 (1912);

Marloth, FI. S. Afr. 4 : 15, fig. 6, t.2 (1915);

Wager in Trans. Roy. Soc. S. Afr. 16 : 191

(1928); Ernst-Schwarzenbach in Ber.

Schweiz. Bot. Ges. 55 : 56 (1945); Oberm. in

Bothalia 8 : 140 (1964). Syntypes: Cape,

Albany, Zeyher; without locality, Drege

(TCD).

Hydril/a dregeana Presl, Bot. Bemerk. 112 (1844).

Type: Cape, Port Elizabeth, Swartkops River,

Drege 2276c. H. muscoides (Harv.) Planch, in Ann.

Sci. Nat. ser. 3, 1 1 : 79 (1849).

Stems 0-5-2 mm in diameter, usually

weak. Leaves alternate, spreading, soft, thin,

transparent, light green, linear, average size

10 mm long and 1 -5 mm broad, apex acute,

margin with 3 rows of hyaline cells which

contain no chlorophyll ; teeth many, acicular,

pointing upwards, leaf-tip with 2 slightly

larger teeth ; cells of lamina elongate-oblong,

with 2-3 rows of cells on each side of the mid-

rib somewhat lighter in colour. Male

inflorescence with spathe-valves flat, obovate,

about 4 mm long and 2 mm broad, with about

23 teeth on each side and with up to 40

flower-buds inside; flowers white to pink.

Female inflorescence with spathe-valves ovoid,

about 4-5 mm long and 1 -5 mm broad, with

about 25 long, acute teeth on each side;

perianth white, maximum length about 25 cm;

ovary flask-shaped, stigmas purple; ovules

about 12, funicles straight (or bent in narrow

ovaries). Capsule narrow-ovoid, 5-10 mm

long, with about 9 seeds breaking away from

the placenta and pushed upwards one behind

the other, each about 2 • 5 mm long. Figs. 31;

32 : 1.

Widespread in the summer rainfall areas of

Southern Africa, from Uitenhage in the eastern

Cape to Natal, Orange Free State, northern Cape,

Transvaal and South West Africa. Also extends

to Bechuanaland, Southern Rhodesia and tropical

east Africa.

Cape. — Albany: near Grahamstown, Dickens s.n.

Barkly West: Holpan, Acocks 2308. Hay: Papkuil-

sloot, Wilman 1247. Port Elizabeth: Gamtoos

River, Schlechter 6059.

O.F.S. — Fauresmith: near Luckhoff, Smith 499.

Kroonstad: Pont 602.

Basutoland. — Banen, Gidllarmod 2066. Leribe:

Makokoane, Dieterlen 1031.

Natal. — Bergville: Edwards 2409.

Transvaal. — Belfast: Obermeyer in TRV 36067.

Benoni: Moss 11175. Christiana: Burtt Davy 12792.

Letaba: Kruger National Park, Bobubu, Van der

Schijff 567 1 . Potchefstroom: Witpoort, Louw 1708.

Pretoria: near Hammanskraal, Mauve & Schlieben

9599. Soutpansberg: Duwenhage’s pan,

Schweickerdt & Verdoorn 627.

S.W.A.— Gibeon: Haribes, Volk 12405. Groot-

fontein: Kumkaus, Kinges 2836; Grosshuis, Schoen-

felder 5684; Naruchas, Dinter 7212; 7384.

Okahandja: Bradfield 390. Rehoboth: Buellsport,

Strey 2118; Rodin 2940.

A form which is much larger in all respects but has

the same distribution as the normal form, is often

found together with it. The following gatherings

belong to it: —

Cape. — Komga: Flanagan 1335.

Natal. — Dundee: Worthington Dam, Hammer .

Estcourt: Tabamhlope Research Station, West 900.

Transvaal. — Petersburg : Blouberg, Strey &

Schlieben 8579. Standerton: Burtt Davy.

S.W.A. — Kaokoveld: Barnard 593.

2. Lagarosiphon verticillifolius Oberm. in

Bothalia 8 ; 142 (1964). Type: Natal, Hlabisa,

Hluhluwe Game Reserve, Ward 2551 $

(PRE, holo.).

Stems 0-5-1 mm in diameter, fairly firm,

densely leafy. Leaves in verticils of 5-7,

occasionally sub-whorled; lamina spreading,

soft, thin, transparent, linear, average size

10 mm (up to 2 cm) long and 0-5 mm broad;.

104

Hydrocharitaceae

Fig. 31. — 1, Lagarosiphon muscoides, X 1 ; a, male floating flower showing reflexed perianth, 3 open stamens

and 3 erect papillate staminodes connate at the tip, x 12; b, female flower with the perianth segments

and 3 forked papillate styles; floating on the surface of the water, x 10; c, young male flowers inside

spathe (diagrammatic), x 7.

Hydrocharitaceae

105

106

Hydrocharitaceae

margin chlorophyllous, teeth many, short,

acicular, pointing out- and upwards, emerging

from a triangular excrescence; with a few

cell-rows beside the midrib slightly larger,

lacunar, the outer row with thickened walls.

Male inflorescence with the spathe-valves

lanceolate, convex, 2-5 mm long and 1 mm

broad, acute, with a few marginal teeth.

Female inflorescence with the spathe-valves

ovoid below, tubular above, apex bifid,

occasionally with a few marginal teeth;

perianth white, stigmatic arms long with large

papillae; ovary with 9-12 ovules, funicle

straight, shorter than the ovule. Capsule

ovoid, 5 mm long and 2 mm broad, elongated

at the apex; seeds 1-5 mm long, narrowly

grooved and with some papillae. Fig. 32 : 2.

Recorded from Natal and Transvaal. Also

found in Mozambique and Rhodesia.

Natal. — Hlabisa: Hluhluwe Game Reserve,

Ward 2518; 2549 <7; 2551 $. Ubombo; Mkuzi

Game Reserve, Tinley 443.

Transvaal. — Letaba: K.N.P., Shangoni,

Mathlambanthlova Pan, Van der Schijff 2857.

Pilgrims Rest: Kruger National Park, 19 miles N.E.

of Skukuza, Codd 6069.

3. Lagarosiphon crispus Rendle in Journ.

Linn. Soc. Bot. 30 : 381, t.31 figs. 8-17 (1895);

Wright in F.T.A. 7 : 4 (1897); Oberm. in

Bothalia 8 : 143 (1964). Type: Tanganyika,

between Uyui and coast, Taylor (K, holo.).

L. tsotsorogensis Brem. & Oberm., in Ann. Transv.

Mus. 14 : 401 (1935). Type: Bechuanaland,

Tsotsoroga Pan, Van Son in TRV 28853 (PRE,

holo.).

Stems filiform, 1-2 mm in diameter.

Leaves alternate, subopposite to verticillate

above, usually densely spaced on the stems,

spreading, soft, thin, transparent, linear,

average size 14 mm long and 0-5 mm broad,

marginal cells chlorophyllous, the many sharp

curved teeth pointing upwards, situated on

small triangular excrescences; leaf-tips bi-

dentate; central area beside the midrib nar-

row to very broad, consisting of large lacunar

cells without or with very little chlorophyll,

usually with distinct transverse veins. Male

inflorescence with spathe-valves cup-shaped,

obovoid, 2 mm long, acute at first, becoming

broad and gaping during anthesis, with about

8 teeth on each side. Female inflorescence

with spathes forming a narrow ovoid tube

2 • 5 mm long, entire or with some small teeth ;

perianth white, limb 2 mm in diameter; ovary

with 24-30 ovules, funicles about as long as

the ovules; stigmas with short papillae.

Capsule ovoid, acuminate, 3 mm long, many-

seeded; seeds 1 mm long, turbinate, with

raised ridges which have transverse septa

(when dry). Fig. 32 : 3; 33.

Recorded from South West Africa and the eastern

and northern Transvaal. Also found in Bechuana-

land, Rhodesia and eastern Africa.

Transvaal. — Pilgrims Rest: Kruger National

Park, Leeupan near Tshokwane, Van der Schijjf

2766; 4198. Soutpansberg: Limpopo River, farm

Weipe, Codd & De Winter 348; Verdoorn 2273.

S.W.A. — Caprivi: Kabuta Village, Chobe River,

Munro. Grootfontein : west of Andara, Merxmuller

2096. Okavango Native Territory: Okavango

River, Niangana, Dinter 7246.

4. Lagarosiphon major ( Ridley ) Moss ex

Wager in Trans. Roy. Soc. S. Afr. 16 : 193

(1928); Mason in New Zeal. Journ. Sci.

30 : 384 (1960); Wild in Kirkia 2 : 39, t,13d

(1961); Oberm. in Bothalia 8 : 145 (1964).

Syntypes: Cape, Victoria East, Umdizine

River, Cooper 17 (K, PRE, BOL); Transvaal,

Bronkhorstspruit, Rehmann 6559 (K, BOL);

Bethal, Trigardtsfontein, Rehmann 6678 (K).

L. muscoides Harv. var. major Ridley in Journ.

Linn. Soc. Bot. 22 : 233 (1886); Wright in F.C.

5, 3 : 2 (1912).

Stems about 3 mm in diameter, usually

firm. Leaves alternate, laxly spaced below,

densely so above, lamina spreading-recurved,

soft but firm, opaque, linear, average size 16

mm (occasionally up to 3 cm) long and 2 mm

broad, apex obtuse or acute to acuminate,

margin with the outermost row of cells

hyaline, forming minute, blunt, closely spaced

teeth; midrib sunk above, prominent below;

lamina with small papillose rhomboid cells,

those of the midrib slightly longer and larger.

Male inflorescence with spathe-valves oblong-

lanceolate, 5 mm long and 3 mm broad, with

about 8 teeth on each side, bearing about 50

flowers consecutively; perianth pinkish. Fe-

male inflorescence with spathe-valves ovoid,

up to 5-5 mm long and 2-5 mm broad, with

about 12 teeth on one side and about 6 on the

other; perianth pale pink, maximum length

15 cm; ovary flask-shaped, stigmas red,

ovules about 12. Capsule ovoid, 5 mm long;

seeds 2 mm long. Fig. 32 : 4.

Widespread in Southern Africa from the eastern

Cape to Natal, Orange Free State and Transvaal.

Also in Rhodesia. Recorded as an adventive in.

Hydrocharitaceae

107

Fig. 33. — Lagarosiphon crispus, part of lamina seen from above showing midrib (m) and net-work of veinlets

(v); ci, large central cells; co, smaller outer cells, X 240.

0-5 mm.

108

Hydrocharitaceae

Europe and New Zealand. Usually sterile. Becomes

a pest at times by choking up dams and rivers.

Cape. — King William’s Town : near King William’s

Town, Sim 1561.

Natal. — Newcastle: west of Laingsnek — Quag-

gasnek road, Edwards 2369.

O.F.S. — Boshoff: Smitskraal, Burtt Davy in

PRE 10731.

Transvaal. — Belfast: Wager in NH49186. Ger-

miston: Birchleigh, Wager in NH49187. Piet

Retief: Kleber s.n. Potchefstroom: Frederikstad,

Louw 1709. Pretoria: Rietvlei Dam, Repton 2001.

Standerton : Schlechter 3464.

5. Lagarosiphon ilicifolius Oberm. in

Bothalia 8:145 (1964). Type: Bechuanaland,

Bridge at Toteng on the N.E. tip of Lake

Ngami, Story 4727 (PRE, holo.).

Stems about 3 mm in diameter (those of

the fertile plants usually more slender), firm.

Leaves alternate, occasionally subopposite,

regularly spaced below with the internodes

3-6 mm long, reflexed or recurved, fairly firm,

opaque, lanceolate, average size 8 mm long

and 5 mm broad (smaller in fertile plants),

apex acute, margin with the outer row of cells

hyaline, with the patent strong teeth situated

on broad triangular excrescences, closely

spaced, leaf-tip bidentate; all cells of the epi-

dermis small, squared and sometimes papil-

lose, midrib faint, depressed, prominent

below, purplish in dried state, its cells some-

what longer and narrower. Male inflorescence

unknown. Female inflorescence with spathe-

valves narrow-ovate, 4 mm long, toothed or

entire, acute, pale wine-red; perianth pale

lilac; ovary ovoid, attenuate, ovules about 9,

funicle erect, short. Capsule ovoid, tapered

above, 5-7 mm long; seeds 2 mm long. Fig.

32 : 5.

Recorded from northern South West Africa.

Also present in northern Bechuanaland, Rhodesia

and Zambia (Zambesi River).

S.W.A. — Okavango Native Territory: Okavango

River, Kapako Camp near Mupini Mission, de

Winter & Marais 4516; Runtu, Merxmiil/er 1890;

Niagana, Dinter 7202a.

95 3. OTTELIA

Ottelia Pers., Synops. PI. 1 : 400 (1805); Benth. & Hook.f., Gen. PI. 3 : 453 (1883); Aschers.

& Guerke in Pflanzenfam. 2, 1 : 257 (1889); Wright in F.T.A. 7 : 6 (1897); Dandy in Journ.

Bot. Lond. 72 : 132 (1934); Phill., Gen. ed. 2 : 58 (1951); Dandy in Hutch., Fam. Flow.

PI. ed. 2, 2 : 541 (1959); den Hartog in FI. Males. 1, 5 : 396 (1957). Type species: O. alismoides

(L.) Pers.

Stratiotes L., Sp. PI. 535 (1753); Gen. PI. ed. 5 : 238 (1754), pro parte, quoad sp. 2.

Damasonium Schreb. in Linn. Gen. PI. ed. 8, 1 : 242 (1789), non Damasonium Mill. (1754).

Boottia Wall., PI. As. Rar. 1 : 51 (1830); Benth. & Hook.f., Gen. PI. 3 : 453 (1883); Aschers. & Guerke,

l.c. 255 (1889); Wright, l.c. 7 (1897); Dandy, l.c. 137 (1934); non Bootia Bigel (1824).

Aquatic, rooted, acaulescent, armed or unarmed annuals in South Africa (rarely

caulescent perennials elsewhere). Roots many, simple. Leaves rosulate submerged or in some

species also with floating blades; lamina of submerged leaves ribbon-like, linear to lanceolate,

the floating blade ovate to cordate; nerves 3-11, distinct, connected by cross-veins, sessile or

petioled, sheathing at the base. Inflorescence pedunculate, dioecious or monoecious and

bisexual; spathes saccate, containing many male flowers but the female and the bisexual

flowers usually solitary in African species; spathes firm, with prominent sometimes muricate

ribs or wings or occasionally smooth; perianth white or yellow in South African species,

tubular, with the 3 outer segments persistent, sepaloid and the 3 inner larger, petaloid, just

exserted above the surface of the water at anthesis, ephemeral, pollination entomophilous

(or flowers often cleistogamous); usually 1-3 male flowers exserted at a time the pedicels

lengthening rapidly at anthesis. Male flowers with 6-15 stamens, anthers bilocular, basifixed.

pollen globular, tuberculate, gynoecium rudimentary. Female flowers with a flask-shaped

ovary with 3-6 parietal placentas intruded towards the centre, styles 3-15, each divided into

2 stigmatic arms, ovules numerous, amphitropous, staminodes present or absent. Fruit

ripening inside the accrescent spathe, the lower part of the perianth-tube forming its beak.

Seeds small, fusiform, tomentose or glabrous.

Hydrocharitaceae

109

About 40 species found in the Palaeotropics with one in Brazil. Four tropical and subtropical species,

endemic in Africa, extending southwards to the northern and eastern parts of Southern Africa ; in permanent

or temporary pools and rivers.

Leaves with an ovate to cordate, floating lamina; plants dioecious 1. O. exserta

Leaves all submerged:

Leaves lanceolate, tapered at the base, margin entire, undulate; unarmed plants (rarely with a few

minute prickles); plants monoecious, flowers bisexual 2. O. ulvifolia

Leaves linear, the margin and also the peduncle minutely toothed; spathe narrow, its wings and ribs

prickly; plants dioecious 3. O. kunenensis

Leaves triangular in cross-section, armed with coarse prickles, which are also present on the peduncle

and spathe; plants dioecious 4. O. muricata

1. Ottelia exserta ( Ridley ) Dandy in

Journ. Bot. Lond. 72 : 137 (1934). Syntypes:

Mozambique, Shire River near Shamo, Kirk;

Shupanga, Kirk; Madagascar, Forbes (BM).

Boottia exserta Ridley in Journ. Linn. Soc. Bot.

22 : 240, t.13 (1886); Wright in F.T.A. 7 : 8 (1897).

B. schinziana Aschers. & Guerke in Pflanzenfam.

2, 1 : 256 (1889), in obs. Type: South West Africa,

Olukonda, Schinz (Bt, Z, BM, PRE, photo).

B. rautanenii Guerke in Festschrift, Aschers., 538

(1904). Syntypes: South West Africa, Ovamboland,

between Olukonda and Unkuambi, Rautanen 49;

50; 97 (Z, PRE, photo).

Ottelia schinziana (Aschers. & Guerke) Dandy

in Journ. Bot. Lond. 72 : 138 (1934).

Dioecious. Leaves: primary leaves

fugacious, submerged, linear to lanceolate,

undulate; later leaves with a petiole which is

triangular in cross-section and a floating,

coriaceous, ovate lamina up to 12-5 cm long

and 5-5 cm broad, cordate, obtuse or

attenuate at the base, apex, rounded, 5-7

nerved. Male inflorescence with a flattened

scape; spathe compressed, smooth, broadly

ovate, 4 cm long and 2-5 cm broad, narrowly

winged; flowers about 60, distichously and

centrifugally arranged, maturing consecutive-

ly with about 1-3 appearing simultaneously;

perianth-tube narrowly cylindrical, exserted

for about 3 cm, outer segments narrowly

lanceolate, 17x7 mm, inner widely obovate,

forming a wide cup, 3x3 cm, white with a

yellowish throat, stamens 12, filaments flat-

tened, scabrid, anthers 15 mm long, narrow,

pistillodes 3, yellow. Female inflorescence with

an angled scape; spathe ovate, 5-5 cm long

and 2-5 cm broad, fleshy, grooved; flower

solitary, with a green perianth-tube, short at

first, lengthening in fruit, outer segments

green, firm, linear-lanceolate, 17 mm long and

5 mm broad, inner obovate, 4 cm long, white,

staminodes 3, filiform, styles 3 short, with the

stigmatic arms longer than the style, flattened,

with the edges densely glandular-fimbriate,

with a globular gland at the base. Fruit nar-

rowly ovoid, 4-5 cm long, beaked. Fig. 34:1.

Recorded from eastern Transvaal and northern

Natal. Also occurs in Madagascar, Malawi,

Rhodesia and Mozambique.

Transvaal. — Pilgrims Rest: Kruger National

Park, pan 19 miles N.E. of Skukuza, Codd 6070;

Leeupan near Tshokwane, Codd 6136. Sibasa:

Wambia, pan, Van der Schijff 3805.

Natal. — Ubombo: Mkuzi Game Reserve, Tinley

616; Ward 3071 .

Measurements given apply to plants seen from

southern Africa; specimens collected in Mozam-

bique and Malawi are far larger in all respects.

2. Ottelia ulvifolia (Planch.) IValp., Ann.

3 : 510 (1852-3); Hutch. & Dalz., F.W.T.A.

2 : 301, fig. 279 (1931); Dandyin J. Bot. Lond.

72 : 138 (1934); Oberm., Schweick. & Ver-

doorn in Bothalia 3 : 225 (1937); Wild in

Rhod. Agric. Journ. 49 : 118 (1952); Kirkia

2 : 34, t. 12b (1961). Syntypes: Madagascar,

Lyall 149; Bojer (K).

Damasonium ulvaefolium Planch, in Ann. Sci.

Nat. ser. 3, 11 : 81 (1849).

Ottelia lancifolia Rich., Tent. FI. Abyss. 2 : 280,

t.95 (1851); Wright in F.T.A. 7 : 7 (1897). Type:

Ethiopia, Shire, Quartin Dillon. O. plantaginea

Welw. ex Rendle in Journ. Linn. Soc. Bot. 22 : 238

(1886); Wright in F.T.A. 7; 7(1897). Type: Angola,

Welwitsch 6469 (BM, holo.). O. vesiculata Ridley

in Journ. Linn. Soc. Bot. 22 : 237 (1886); Wright

in F.T.A. 7 : 7 (1897). Type: Angola, Welwitsch

6497 (BM, holo.). O. crassifolia (Ridley) Welw. ex

Rendle in Cat. Afr. PI. Welw. 2, 1 : 2 (1899).

O. baumii Guerke in Baum, Kunene-Sambesi

Exp. 171 (1903). Type: Angola, Baum 858 (B,

holo., M, PRE, photo.). O. benguellensis Guerke in

Baum, Kunene-Sambesi Exp. 172 (1903). Type:

Angola, Lion pan, Baum 120 (B, holo.?). “O. sp.

cf. O. ulvaefolia (Planch.) Walp.”, Wager in Trans.

Roy, Soc. S. Afr. 16 : 201, t.22 (1928). O. australis

Brem. in Ann. Transv. Mus. 15 : 235 (1933). Type:

Transvaal, Pietersburg district, Vivo vlei, Breme-

kamp & Schweickerdt 203 (PRE, holo.). O.

macrantha (Wright) Dandy in Journ. Bot. Lond.

72 : 138 (1934). O. vernayi Brem. & Oberm. in

110

Hydrocharitaceae

Fig. 34.— 1, Ottella exserta, male, x i; la, female flower showing spathe,

inner perianth segment (cut off), one staminode with basal gland and

in florescence; 2, O. kunenensis, female, X

one outer perianth segment, one

one forked style, x 1; lb, male

Hydrocharitaceae

111

Ann. Transv. Mus. 16 : 40! (1935). Type: Bechuana-

land, Chobe River, Maun, Van Son in TRV 28852

(PRE, holo.).

Boottia abyssinica Ridley in Journ. Linn. Soc.

Bot. 22 : 239 (1886); Wright in F.T.A. 7 : 9 (1897).

Type: Ethiopia, Schimper 1452 (BM, holo.).

B. crassifolia Ridley in Journ. Linn. Soc. Bot.

22 : 239 (1886); Wright in F.T.A. 7 : 9 (1897).

Type: Angola, Mumpulla River and at Lopollo,

Welwitsch 6470 (BM, holo.; M, PRE, photo.).

B. rohrbachiana Aschers. & Guerke in Pflanzenfam.

2, 1 : 256 (1889). Type: Central Africa. B. macrantha

Wright in F.T.A. 7 : 9 (1897). Type: Mozambique,

Scott (K, holo.).

Monoecious. Leaves all submerged, in

rosettes, crowded, radical or on petioles, thin

but firm, variable in size and shape, linear-

lanceolate to ovate-lanceolate, up to 23 cm

long and 5 cm broad, attenuate above and

below, margin wavy, entire or sparsely denti-

culate. Inflorescence on a peduncle which is

triangular in cross-section, up to 23 cm long

and 3 mm in diam.; spathes saccate, com-

pressed, narrow-ovate in outline, becoming

much broader in fruit, up to 4 cm long and 2

cm broad, green with brownish nerves, nar-

rowly winged, orifice with a few irregular

small teeth; flowers bisexual; perianth-tube

just exserted from spathe, outer segments

linear-oblong, 8 mm long and 2 mm broad,

green, inner obovate, 25 mm long and 9 mm

broad, yellow or cream, with a small gland at

the base ; stamens 6, filaments flattened, 4 mm

long, anthers small; ovary 17 mm long and

4 mm broad, placentas 6, unequal, styles 6

with 2 short stigmatic arms. Fruit enclosed in

the spathe, seeds small, cylindrical, covered

with fine white hairs. Fig. 35 : 1.

Recorded from the warmer parts of the Trans-

vaal and northern South West Africa. Widespread

in subtropical and tropical Africa.

Transvaal. — Letaba: Gravelotte, Blackhills,

Wager ; Kruger National Park, Mahlambanhlovu

Pan, Van der Schijff 2847. Pietersburg: Vivo vlei,

Bremekamp & Schweickerdt 203. Pretoria: near

Petronella, Dyer, Verdoorn & Mauve 4267.

Soutpansberg: Duvenagespan near Amisfort,

Schweickerdt & Verdoorn 623. Waterberg: near

Naboomspruit, Nyl River, Mauve 4260.

S.W.A. — Caprivi: near Andara, Okavango River,

Merxmidler 2095. Okavango: near Runtu,

Omuramba Omatako, De Winter <£ Marais 5045;

Kanovlei, Dinter 7191.

3. Ottelia kunenensis ( Guerke ) Dandy in

Journ. Bot. Lond. 72 : 137 (1934). Type:

Angola, Kunene River, between Kiteve and

Humbe, Baum 962 (B| holo, Z, PRE, photo.).

Boottia kunenensis Guerke in Baum, Kunene-

Sambesi Exped. 172 (1903).

Dioecious. Leaves submerged, ribbon-

like, linear, up to 50 cm long and 2 cm broad,

indistinctly 5-7 nerved, often with purple

blotches, margin denticulate, undulate, midrib

dorsally denticulate. Inflorescence on a

slender scape, triangular in cross-section,

serrulate or smooth, in the female tightly

coiled in fruit ; male flower with spathe ovoid,

up to 3 cm long and 1 cm broad, with 9-12

narrow denticulate ribs, orifice split into 3-9

teeth, outer segments linear, 14 mm long and

2 mm broad, inner obovate, about 25 mm

long, white or yellow, stamens 12-15 of

various lengths, rudimentary ovary glandular,

3-lobed; female spathe cylindrical, 2-5 cm

long, perianth-tube as long as the spathe,

segments resembling those of the male, the 6

styles divided nearly to the base with the

stigmatic arms about 15 mm long. Fig. 34 : 2.

Recorded from South West Africa. Also in

Angola.

S.W.A. — Okavango Native Territory: Okavango

River at Runtu, De Winter & Marais 4909 ; Merx-

midler 1882; Niangana, Dinter 7250. Caprivi:

Kabuta Village, Chobe River, Munro ML3.

4. Ottelia muricata ( Wright ) Dandy in

Journ. Bot. Lond. 72 : 138 (1934). Syntypes:

Bechuanaland, Botletle River, Lugard 13;

Chobe River, McCabe 32 (K).

Boottia muricata Wright in F.T.A. 7 : 569 (1898);

Hook., Ic. PL t.3 101 (1933), partly, excl. fig. 1.

B. aschersoniana Guerke in Baum, Kunene-Sambesi

Exped. 173 (1903); Dandy in Journ. Bot. Lond.

72 : 138 (1934). Type: Angola, Kuito depression

between Kutui and Sobi, Baum 772 (B).

Dioecious, perennial, armed (the conical

projections soft in the living state). Leaves

submerged, filiform, with a dorsal prominent

rib, to triangular with short coarse teeth on

the ribs, lamina up to 2 m long and 5-15 mm

wide. Inflorescence with the spinous scape

terete to triangular in cross-section, resem-

bling the leaves; male flower with spathe

fusiform, about 35 mm long and 10 mm

broad, leathery, with 7-10 spiny ribs, orifice

bilobed, perianth-tube exserted for 14-30

mm, outer segments linear-lanceolate, 14 mm

long, inner 2-3 cm long, obovate, white with

an orange throat, minutely papillate outside,

stamens 12, of various lengths; female flower

with spathe cylindrical, 45 mm long and 4 mm

broad, with about 7 spiny ribs, perianth-tube

112

Hydrocharitaceae

exserted for about 7 mm from the spathe,

styles 3, short, each with 2 stigmatic arms

12 mm long; staminodes 6, three with long

papillate filaments, 3 small, scale-like. Fig.